Climate and sea-level variations along the northwestern Tethyan margin during the Valanginian C-isotope excursion: Mineralogical evidence from the Vocontian Basin (SE France)

A high resolution mineralogical study (bulk-rock and clay-fraction) was carried out upon the hemipelagic strata of the Angles section (Vocontian Basin, SE France) in which the Valanginian positive C-isotope excursion occurs. To investigate sea-level fluctuations and climate change respectively, a Detrital Index (DI: (phyllosilicates and quartz)/calcite) and a Weathering Index (WI: kaolinite/(illite + chlorite)) were established and compared to second-order sea-level fluctuations. In addition, the mineralogical data were compared with the High Nutrient Index (HNI, based on calcareous nannofossil taxa) data obtained by Duchamp-Alphonse et al. (2007), in order to assess the link between the hydrolysis conditions recorded on the surrounding continents and the trophic conditions inferred for the Vocontian Basin. It appears that the mineralogical distribution along the northwestern Tethyan margin is mainly influenced by sea-level changes during the Early Valanginian (Pertransiens to Stephanophorus ammonite Zones) and by climate variations from the late Early Valanginian to the base of the Hauterivian (top of the Stephanophorus to the Radiatus ammonite Zones). The sea-level fall observed in the Pertransiens ammonite Zone (Early Valanginian) is well expressed by an increase in detrital inputs (an increase in the DI) associated with a more proximal source and a shallower marine environment, whereas the sea-level rise recorded in the Stephanophorus ammonite Zone corresponds to a decrease in detrital influx (a decrease in the DI) as the source becomes more distal and the environment deeper. Interpretation of both DI and WI, indicates that the positive C-isotope excursion (top of the Stephanophorus to the Verrucosum ammonite Zones) is associated with an increase of detrital inputs under a stable, warm and humid climate, probably related to greenhouse conditions, the strongest hydrolysis conditions being reached at the maximum of the positive C-isotope excursion. From the Verrucosum ammonite Zone to the base of the Hauterivian (Radiatus ammonite Zone) climatic conditions evolved from weak hydrolysis conditions and, most likely, a cooler climate (resulting in a decrease in detrital inputs) to a seasonal climate in which more humid seasons alternated with more arid ones. The comparison of the WI to the HNI shows that the nutrification recorded al: the Angles section from the top of the Stephanophorus to the Radiatus ammonite Zones (including the positive C-isotope shift), is associated with climatic changes in the source areas. At that time, increased nutrient inputs were generally triggered by increased weathering processes in the source areas due to acceleration in the hydrological cycle under greenhouse conditions This scenario accords with the widely questioned palaeoenvironmental model proposed by Lini et al., (1992) and suggests that increasing greenhouse conditions are the main factor that drove the palaeoenvironmental changes observed in the hemipelagic realm of the Vocontian Basin, during the Valanginian positive C-isotope shift. This high-resolution mineralogical study highlights short-term climatic changes during the Valanginian, probably associated to rapid changes in the C-cycle. Coeval Massive Parana-Etendeka flood basalt eruptions may explain such rapid perturbations. (C) 2011 Elsevier B.V. All rights reserved.

Growth and demise of Permian biogenic chert along northwest Pangea: evidence for end-Permian collapse of thermohaline circulation

The Permian Chert Event (PCE) was a 30 Ma long episode of unusual chert accumulation along the northwest margin of Pangea, and possibly worldwide. The onset of the PCE occurred at about the Sakmarian-Artinskian boundary in the Sverdrup Basin, Canadian Arctic, where it coincides with a maximum flooding event, the ending of high-frequency/high-amplitude shelf cyclicity, the onset of massive biogenic chert deposition in deep-water distal areas, and a long-term shift from warm- to cool-water carbonate sedimentation in shallow-water proximal areas. A similar and coeval shift is observed from the Barents Sea to the northwestern USA. A landward and southward expansion of silica factories occurred during the Middle and Late Permian at which time warm-water carbonate producers disappeared completely from the northwest margin of Pangea. Biotically impoverished and increasingly narrow cold-water carbonate factories (characterised by non-cemented bioclasts of sponges, bryozoans, echinoderms and brachiopods) were then progressively replaced by silica factories. By Late Permian time, little carbonate sediments accumulated in the Barents Sea and in the Sverdrup Basin. where the deep- to shallow-water sedimentary spectrum was occupied by siliceous sponge spicules. By that time, biogenic silica sedimentation was common throughout the world. Silica factories collapsed in the Late Permian, abruptly bringing the PCE to an end. In northwest Pangea, the end- Permian collapse of the PCE was associated with a major transgression and with a return to much warmer oceanic and continental climatic conditions. Chert deposition resumed in the distal oceanic areas during the early Middle Triassic (Anisian) after a 8-10 Ma interruption (Early Triassic Chert Gap). The conditions necessary for the onset, expansion and zenith of the PCE were provided by the thermohaline circulation of nutrient-rich cold waters along the northwestern and western margin of Pangea, and possibly throughout the world oceans. These conditions provided an efficient transportation mechanism that constantly replenished the supply of silica in the area, created a nutrient- and oxygen-rich environment favouring siliceous biogenic productivity. established cold sea-floor conditions, hindering silica dissolution, while increasing calcium carbonate solubility, and provided conditions adverse to organic and inorganic carbonate production, The northwest margin of Pangea was, for nearly 30 Ma. bathed by cold waters presumably derived from the seasonal melting of northern sea ice, the assumed engine for thermohaline circulation. This process started near the Sakmarian-Artinskian boundary. intensified throughout Middle and Late Permian time and ceased suddenly in latest Permian time, It led to oceanic conditions much colder than normally expected from the palaeolatitudes. and the influence of cold northerly-derived water was felt as far south southern Nevada. The demise of silica factories was caused by the rapid breakdown of these conditions and the establishment of a much warmer marine environment accompanied by sluggish circulation and perhaps a reduced input of dissolved silica to the ocean. Complete thawing of northern sea ice would have ended thermohaline circulation and led to warm and sluggish oceanic conditions inimical to the production. accumulation and preservation of biogenic silica.

Community replacement of neritic carbonate organisms during the late Valanginian platform demise: A new record from the Provence Platform

The Valanginian is marked by a major platform demise inducing a hiatus in the northern Tethyan neritic carbonate record from the top of the lower Valanginian to the lower Hauterivian. New biostratigraphic and chemostratigraphic data from the Ollioules section (Provence Platform, southern France) are presented here, demonstrating that a large part of the upper Valanginian is preserved in an inner platform environment. The thick, upper Valanginian, aggrading carbonate succession is observed in an aborted rift domain, implying relatively low subsidence. In this context, a relatively long-term sea-level rise was required to sustain a keep-up style of carbonate production. Like the Apulian Platform, the remarkable preservation of the Provence Platform may have been favored by its remoteness from terrigenous source areas, as suggested by the low clastic inputs and low P-accumulation rates. Two main biotic community replacements are observed in Ollioules. The first saw the development of abundant microbialites and algae at the onset of the late Valanginian. A Tubiphytes concentration occurred during the coolest climatic conditions and the transition towards arid conditions, whereas the subsequent Lithocodium-Bacinella and orbitolinids assemblages developed under low nutrient conditions during a warmer interval. Both assemblages may have been triggered by increased alkalinity. The second community replacement saw the installation of coral- and rudist-dominated communities during the latest Valanginian to early Hauterivian. They indicate a change to oligotrophic, open marine conditions. Six medium-scale sequences have been defined in Ollioules, indicating short-term transgressive-regressive trends superimposed on a long-term transgression. Low nutrient inputs and relatively low subsidence in an aggradational context may explain the survival of the isolated Provence Carbonate Platform during a time of widespread drowning episodes and platform demise in the northern Tethyan domain. (c) 2012 Elsevier B.V. All rights reserved.

Introgressive hybridization of threatened European tree frogs (Hyla arborea) by introduced H. intermedia in Western Switzerland

Hybridization by introduced taxa is a major threat to native species. Characterizing human introductions is thus one of the missions of conservation geneticists. Here we survey a declining population of the regionally endangered European tree frog (Hyla arborea) in the Grangettes natural reserve (Rhone valley, Western Switzerland), where previous evidence indicated human introduction of the Italian taxon H. intermedia. We combined fast-evolving mitochondrial and nuclear markers and an extended sampling to conduct population genetic analyses of the Grangettes and putative source areas. We show that the Grangettes population is a hybrid swarm, with all individuals featuring recent nuclear admixture and mitochondrial DNA of introduced H. intermedia, most likely of proximate south Alpine origin. In contrast, H. arborea and H. intermedia hardly introgress in their natural parapatric ranges, consistent with an advanced reproductive isolation. Thus, potential hybrid incompatibilities may account for the strong decline of this population, despite important conservation efforts. Although their hybrid nature makes them a priori unworthy of any protection, we propose specific measures to recover local H. arborea gene pool and preserve tree frogs in the Grangettes, the last population remaining from this heavily impacted part of the Alps.

Rockfall susceptibility assessment and remote geological mapping with LiDAR point clouds

Characterizing the geological features and structures in three dimensions over inaccessible rock cliffs is needed to assess natural hazards such as rockfalls and rockslides and also to perform investigations aimed at mapping geological contacts and building stratigraphy and fold models. Indeed, the detailed 3D data, such as LiDAR point clouds, allow to study accurately the hazard processes and the structure of geologic features, in particular in vertical and overhanging rock slopes. Thus, 3D geological models have a great potential of being applied to a wide range of geological investigations both in research and applied geology projects, such as mines, tunnels and reservoirs. Recent development of ground-based remote sensing techniques (LiDAR, photogrammetry and multispectral / hyperspectral images) are revolutionizing the acquisition of morphological and geological information. As a consequence, there is a great potential for improving the modeling of geological bodies as well as failure mechanisms and stability conditions by integrating detailed remote data. During the past ten years several large rockfall events occurred along important transportation corridors where millions of people travel every year (Switzerland: Gotthard motorway and railway; Canada: Sea to sky highway between Vancouver and Whistler). These events show that there is still a lack of knowledge concerning the detection of potential rockfalls, making mountain residential settlements and roads highly risky. It is necessary to understand the main factors that destabilize rocky outcrops even if inventories are lacking and if no clear morphological evidences of rockfall activity are observed. In order to increase the possibilities of forecasting potential future landslides, it is crucial to understand the evolution of rock slope stability. Defining the areas theoretically most prone to rockfalls can be particularly useful to simulate trajectory profiles and to generate hazard maps, which are the basis for land use planning in mountainous regions. The most important questions to address in order to assess rockfall hazard are: Where are the most probable sources for future rockfalls located? What are the frequencies of occurrence of these rockfalls? I characterized the fracturing patterns in the field and with LiDAR point clouds. Afterwards, I developed a model to compute the failure mechanisms on terrestrial point clouds in order to assess the susceptibility to rockfalls at the cliff scale. Similar procedures were already available to evaluate the susceptibility to rockfalls based on aerial digital elevation models. This new model gives the possibility to detect the most susceptible rockfall sources with unprecented detail in the vertical and overhanging areas. The results of the computation of the most probable rockfall source areas in granitic cliffs of Yosemite Valley and Mont-Blanc massif were then compared to the inventoried rockfall events to validate the calculation methods. Yosemite Valley was chosen as a test area because it has a particularly strong rockfall activity (about one rockfall every week) which leads to a high rockfall hazard. The west face of the Dru was also chosen for the relevant rockfall activity and especially because it was affected by some of the largest rockfalls that occurred in the Alps during the last 10 years. Moreover, both areas were suitable because of their huge vertical and overhanging cliffs that are difficult to study with classical methods. Limit equilibrium models have been applied to several case studies to evaluate the effects of different parameters on the stability of rockslope areas. The impact of the degradation of rockbridges on the stability of large compartments in the west face of the Dru was assessed using finite element modeling. In particular I conducted a back-analysis of the large rockfall event of 2005 (265'000 m3) by integrating field observations of joint conditions, characteristics of fracturing pattern and results of geomechanical tests on the intact rock. These analyses improved our understanding of the factors that influence the stability of rock compartments and were used to define the most probable future rockfall volumes at the Dru. Terrestrial laser scanning point clouds were also successfully employed to perform geological mapping in 3D, using the intensity of the backscattered signal. Another technique to obtain vertical geological maps is combining triangulated TLS mesh with 2D geological maps. At El Capitan (Yosemite Valley) we built a georeferenced vertical map of the main plutonio rocks that was used to investigate the reasons for preferential rockwall retreat rate. Additional efforts to characterize the erosion rate were made at Monte Generoso (Ticino, southern Switzerland) where I attempted to improve the estimation of long term erosion by taking into account also the volumes of the unstable rock compartments. Eventually, the following points summarize the main out puts of my research: The new model to compute the failure mechanisms and the rockfall susceptibility with 3D point clouds allows to define accurately the most probable rockfall source areas at the cliff scale. The analysis of the rockbridges at the Dru shows the potential of integrating detailed measurements of the fractures in geomechanical models of rockmass stability. The correction of the LiDAR intensity signal gives the possibility to classify a point cloud according to the rock type and then use this information to model complex geologic structures. The integration of these results, on rockmass fracturing and composition, with existing methods can improve rockfall hazard assessments and enhance the interpretation of the evolution of steep rockslopes. -- La caractérisation de la géologie en 3D pour des parois rocheuses inaccessibles est une étape nécessaire pour évaluer les dangers naturels tels que chutes de blocs et glissements rocheux, mais aussi pour réaliser des modèles stratigraphiques ou de structures plissées. Les modèles géologiques 3D ont un grand potentiel pour être appliqués dans une vaste gamme de travaux géologiques dans le domaine de la recherche, mais aussi dans des projets appliqués comme les mines, les tunnels ou les réservoirs. Les développements récents des outils de télédétection terrestre (LiDAR, photogrammétrie et imagerie multispectrale / hyperspectrale) sont en train de révolutionner l'acquisition d'informations géomorphologiques et géologiques. Par conséquence, il y a un grand potentiel d'amélioration pour la modélisation d'objets géologiques, ainsi que des mécanismes de rupture et des conditions de stabilité, en intégrant des données détaillées acquises à distance. Pour augmenter les possibilités de prévoir les éboulements futurs, il est fondamental de comprendre l'évolution actuelle de la stabilité des parois rocheuses. Définir les zones qui sont théoriquement plus propices aux chutes de blocs peut être très utile pour simuler les trajectoires de propagation des blocs et pour réaliser des cartes de danger, qui constituent la base de l'aménagement du territoire dans les régions de montagne. Les questions plus importantes à résoudre pour estimer le danger de chutes de blocs sont : Où se situent les sources plus probables pour les chutes de blocs et éboulement futurs ? Avec quelle fréquence vont se produire ces événements ? Donc, j'ai caractérisé les réseaux de fractures sur le terrain et avec des nuages de points LiDAR. Ensuite, j'ai développé un modèle pour calculer les mécanismes de rupture directement sur les nuages de points pour pouvoir évaluer la susceptibilité au déclenchement de chutes de blocs à l'échelle de la paroi. Les zones sources de chutes de blocs les plus probables dans les parois granitiques de la vallée de Yosemite et du massif du Mont-Blanc ont été calculées et ensuite comparés aux inventaires des événements pour vérifier les méthodes. Des modèles d'équilibre limite ont été appliqués à plusieurs cas d'études pour évaluer les effets de différents paramètres sur la stabilité des parois. L'impact de la dégradation des ponts rocheux sur la stabilité de grands compartiments de roche dans la paroi ouest du Petit Dru a été évalué en utilisant la modélisation par éléments finis. En particulier j'ai analysé le grand éboulement de 2005 (265'000 m3), qui a emporté l'entier du pilier sud-ouest. Dans le modèle j'ai intégré des observations des conditions des joints, les caractéristiques du réseau de fractures et les résultats de tests géoméchaniques sur la roche intacte. Ces analyses ont amélioré l'estimation des paramètres qui influencent la stabilité des compartiments rocheux et ont servi pour définir des volumes probables pour des éboulements futurs. Les nuages de points obtenus avec le scanner laser terrestre ont été utilisés avec succès aussi pour produire des cartes géologiques en 3D, en utilisant l'intensité du signal réfléchi. Une autre technique pour obtenir des cartes géologiques des zones verticales consiste à combiner un maillage LiDAR avec une carte géologique en 2D. A El Capitan (Yosemite Valley) nous avons pu géoréferencer une carte verticale des principales roches plutoniques que j'ai utilisé ensuite pour étudier les raisons d'une érosion préférentielle de certaines zones de la paroi. D'autres efforts pour quantifier le taux d'érosion ont été effectués au Monte Generoso (Ticino, Suisse) où j'ai essayé d'améliorer l'estimation de l'érosion au long terme en prenant en compte les volumes des compartiments rocheux instables. L'intégration de ces résultats, sur la fracturation et la composition de l'amas rocheux, avec les méthodes existantes permet d'améliorer la prise en compte de l'aléa chute de pierres et éboulements et augmente les possibilités d'interprétation de l'évolution des parois rocheuses.

Aptian to Cenomanian deeper-water hiatal stromatolites from the northern Tethyan margin

A suite of deeper-water hiatal (DWH) stromatolites has been identified in the phosphatic and glauconitic sediments of Aptian to Cenomanian age in the alpine Helvetic thrust-and-fold belt, which represents the former northern Tethyan margin. The most important occurrences date from the latest Early to Late Aptian, the late Early to early middle Albian, and the Early Cenomanian. They are invariably associated with condensed phosphatic beds and occur preferentially on top of hardgrounds or on reworked pebbles and fossils. The zone of optimal stromatolite growth and preservation coincides with the zone of maximal sedimentary condensation, in the deeper parts of phosphogenic areas. The DWH stromatolites show variable morphologies, ranging from isolated laminae ("films") to internally laminated columns and crusts. They reach thicknesses of maximal 10 cm and are either preserved in phosphate or micrite. In the latter case, they may show peripheral impregnations of phosphate or iron oxyhydroxides. The quasi-complete lack of macroscopic sessile organisms suggests that the DWH stromatolites grew close to the upper boundary of an oxygen-minimum zone. Electron-scanning microscopic images show that the Early Cenomanian examples preserved in micrite consist of filamentous structures, which form spaghetti-like assemblages. They are. interpreted as the remains of poikiloaerobic, heterotrophic microbes. Coeval DWH stromatolites are known from the entire European segment of the northern Tethyan margin, and shallow-water counterparts are commonplace on Tethyan carbonate platforms. This indicates that, in general, paleoceanographic and paleoenvironmental conditions were appropriate for stromatolite growth and preservation. The here-described DWH stromatolites proliferated especially in time windows, which followed upon the oceanic anoxic periods OAE la (Early Aptian), lb (latest Aptian and earliest Albian), and Id (latest Albian). They may represent pioneer ecosystems, which thrived during the recovery phases following the "mid"-Cretaceous OAEs.

L'exploitation des eaux souterraines en Syrie centrale : construction de diagnostics et politiques d'intervention

Syrian dry areas have been for several millennia a place of interaction between human populations and the environment. If environmental constraints and heterogeneity condition the human occupation and exploitation of resources, socio-political, economic and historical elements play a fundamental role. Since the late 1980s, Syrian dry areas are viewed as suffering a serious water crisis, due to groundwater overdraft. The Syrian administration and international development agencies believe that groundwater overexploitation is also leading to a decline of agricultural activities and to poverty increase. Action is thus required to address these problems.However, the overexploitation diagnosis needs to be reviewed. The overexploitation discourse appears in the context of Syria's opening to international organizations and to the market economy. It echoes the international discourse of "global water crisis". The diagnosis is based on national indicators recycling old Soviet data that has not been updated. In the post-Soviet era, the Syrian national water policy seems to abandon large surface water irrigation projects in favor of a strategy of water use rationalization and groundwater conservation in crisis regions, especially in the district of Salamieh.This groundwater conservation policy has a number of inconsistencies. It is justified for the administration and also probably for international donors, since it responds to an indisputable environmental emergency. However, efforts to conserve water are anecdotal or even counterproductive. The water conservation policy appears a posteriori as an extension of the national policy of food self-sufficiency. The dominant interpretation of overexploitation, and more generally of the water crisis, prevents any controversary approach of the status of resources and of the agricultural system in general and thus destroys any attempt to discuss alternatives with respect to groundwater management, allocation, and their inclusion in development programs.A revisited diagnosis of the situation needs to take into account spatial and temporal dimensions of the groundwater exploitation and to analyze the co-evolution of hydrogeological and agricultural systems. It should highlight the adjustments adopted to cope with environmental and economic variability, changes of water availability and regulatory measures enforcements. These elements play an important role for water availability and for the spatial, temporal, sectoral allocation of water resource. The groundwater exploitation in the last century has obviously had an impact on the environment, but the changes are not necessarily catastrophic.The current groundwater use in central Syria increases the uncertainty by reducing the ability of aquifers to buffer climatic changes. However, the climatic factor is not the only source of uncertainty. The high volatility of commodity prices, fuel, land and water, depending on the market but also on the will (and capacity) of the Syrian State to preserve social peace is a strong source of uncertainty. The research should consider the whole range of possibilities and propose alternatives that take into consideration the risks they imply for the water users, the political will to support or not the local access to water - thus involving a redefinition of the economic and social objectives - and finally the ability of international organizations to reconsider pre-established diagnoses.

Sr, C and O isotope systematics in the Pucara Basin, central Peru - Comparison between Mississippi Valley-type deposits and barren areas

A combined Sr, O and C isotope study has been carried out in the Pucara basin, central Peru, to compare local isotopic trends of the San Vicente and Shalipayco Zn-Pb Mississippi Valley-type (MVT) deposits with regional geochemical patterns of the sedimentary host basin. Gypsum, limestone and regional replacement dolomite yield Sr-87/Sr-86 ratios that fall within or slightly below the published range of seawater Sr-87/Sr-86 values for the Lower Jurassic and the Upper Triassic. Our data indicate that the Sr isotopic composition of seawater between the Hettangian and the Toarcian may extend to lower Sr-87/Sr-86 ratios than previously published values. An Sr-87-enrichment is noted in (1) carbonate rocks from the lowermost part of the Pucara basin, and (2) different carbonate generations at the MVT deposits. This indicates that host rocks at MVT deposits and in the lowermost part of the carbonate sequence interacted with Sr-87-enriched fluids. The fluids acquired their radiogenic nature by interaction with lithologies underlying the carbonate rocks of the Pucara basin. The San Ramon granite, similar Permo-Triassic intrusions and their elastic derivatives in the Mitu Group are likely sources of radiogenic Sr-87. The Brazilian shield and its erosion products are an additional potential source of radiogenic Sr-87. Volcanic rocks of the Mitu Group are not a significant source for radiogenic Sr-87; however, molasse-type sedimentary rocks and volcaniclastic rocks cannot be ruled out as a possible source of radiogenic Sr-87. The marked enrichment in Sr-87 of carbonates toward the lower part of the Pucara Group is accompanied by only a slight decrease in delta(18)O values and essentially no change in delta(13)C values, whereas replacement dolomite and sparry carbonates at the MVT deposits display a coherent trend of progressive Sr-87-enrichment, and O-18- and C-13-depletion. The depletion in O-18 in carbonates from the MVT deposits are likely related to a temperature increase, possibly coupled with a O-18-enrichment of the ore-forming fluids. Progressively lower delta(13)C values throughout the paragenetic sequence at the MVT deposits are interpreted as a gradually more important contribution from organically derived carbon. Quantitative calculations show that a single fluid-rock interaction model satisfactorily reproduces the marked Sr-87-enrichment and the slight decrease in delta(18)O values in carbonate rocks from the lower part of the Pucara Group. By contrast, the isotopic covariation trends of the MVT deposits are better reproduced by a model combining fluid mixing and fluid-rock interaction. The modelled ore-bearing fluids have a range of compositions between a hot, saline, radiogenic brine that had interacted with lithologies underlying the Pucara sequence and cooler, dilute brines possibly representing local fluids within the Pucara sequence. The composition of the local fluids varies according to the nature of the lithologies present in the neighborhood of the different MVT deposits. The proportion of the radiogenic fluid in the modelled fluid mixtures interacting with the carbonate host rocks at the MVT deposits decreases as one moves up in the stratigraphic sequence of the Pucara Group.

Sulfur and strontium isotope composition of the Llobregat River (NE Spain): Tracers of natural and anthropogenic chemicals in stream waters

The use of sulfur and strontium isotopes as tracers for the source/s of water contaminants have been applied to the water of the Llobregat River system (NE Spain). Surface water samples from June 1997 were collected from the Llobregat River and its main tributaries and creeks. The chemistry of most stream waters are controlled mainly by the weathering of Tertiary chemical sediments within the drainage basin. The largest variation in delta(34)S values were found in the small creeks with values ranging from -9.9 to 15parts per thousand, whilst in the main river channels values ranged from 6.3 to 12.4parts per thousand. The Sr-87/Sr-86 ratio for dissolved strontium ranged from 0.70795 for a non-polluted site to 0.70882 for a polluted one. Most of the waters with high NO3 and low Ca/Na ratio converge to the same Sr-87/Sr-86 value, pointing to dominant pollutant end member contribution or a mixing of pollutants with an isotopic composition around 0.7083-0.7085. Although the concentration of the natural inputs in the river for sulfate and strontium are high, as a result of the sulfate outcrops within the geology of the basin, their isotopic characteristics suggest that they can be used as a discriminating device in water pollution problems. However to establish the detailed characteristics of the isotopes as geochemical tools, specific high-resolution case studies are necessary in small areas, where the inputs are well known.

Sulfur speciation and stable isotope trends of water-soluble sulfates in mine tailings profiles

Sulfur speciation and the sources of water-soluble sulfate in three oxidizing sulfidic mine tailings impoundments were investigated by selective dissolution and stable isotopes. The studied tailings impoundments-Piuquenes, Cauquenes, and Salvador No. 1-formed from the exploitation of the Rio Blanco/La Andina, El Teniente, and El Salvador Chilean porphyry copper deposits, which are located in Alpine, Mediterranean, and hyperarid climates, respectively. The water-soluble sulfate may originate from dissolution of primary ore sulfates (e.g., gypsum, anhydrite, jarosite) or from oxidation of sulfide minerals exposed to aerobic conditions during mining activity. With increasing aridity and decreasing pyrite content of the tailings, the sulfur speciation in the unsaturated oxidation zones showed a trend from dominantly Fe(Ill) oxyhydroxide fixed sulfate (e.g., jarosite and schwertmannite) in Piuquenes toward increasing presence of water-soluble sulfate at Cauquenes and Salvador No. 1. In the saturated primary zones, sulfate is predominantly present in water-soluble form (mainly as anhydrite and/or gypsum). In the unsaturated zone at Piuquenes and Cauquenes,the delta(34)S(S04) values ranged from +0.5 parts per thousand to +2.0 parts per thousand and from -0.4 parts per thousand to +1.4 parts per thousand Vienna Canyon Diablo Troilite (V-CDT), respectively, indicating a major sulfate source from pyrite oxidation (delta(34)S(pyrite) -1.1 parts per thousand and -0.9 parts per thousand). In the saturated zone at Piuquenes and Cauquenes, the values ranged from -0.8%. to +0.3 parts per thousand and from +2.2 parts per thousand to +3.9 parts per thousand, respectively. At Cauquenes the 34 S enrichment in the saturated zone toward depth indicates the increasing contribution of isotopically heavy dissolved sulfate from primary anhydrite (similar to+10.9%o). At El Salvador No. 1, the delta(34)S(SO4) average value is -0.9 parts per thousand, suggesting dissolution of supergene sulfate minerals (jarosite, alunite, gypsum) with a delta(34)S similar to -0.7 parts per thousand as the most probable sulfate source. The gradual decrease Of delta(18)O(S04) values from the surface to the oxidation front in the tailings impoundments at Piuquenes (from -4.5 parts per thousand to -8.6 parts per thousand Vienna Standard Mean Ocean Water, V-SMOW) and at Cauquenes (from -1.3 parts per thousand to -3.5 parts per thousand) indicates the increasing importance of ferric iron as the main electron acceptor in the oxidation of pyrite. The different delta(18)O(SO4) values between the tailings impoundments studied here reflect the local climates.

Remediation of a marine shore tailings deposit and the importance of water-rock interaction on element cycling in the coastal aquifer

We present the study of the geochemical processes associated with the first successful remediation of a marine shore tailings deposit in a coastal desert environment (Bahia de Ite, in the Atacama Desert of Peru). The remediation approach implemented a wetland on top of the oxidized tailings. The site is characterized by a high hydrauliz gradient produced by agricultural irrigation on upstream gravel terraces that pushed river water (similar to 500 mg/L SO(4)) toward the sea and through the tailings deposit. The geochemical and isotopic (delta(2)H(water) and delta(18)O(water), delta(34)S(sulfate) , delta(18)O(sulfate)) approach applied here revealed that evaporite horizons (anhydrite and halite) in the gravel terraces are the source of increased concentrations of SO(4), Cl, and Na up to similar to 1500 mg/L in the springs at the base of the gravel terraces. Deeper groundwater interacting with underlying marine sequences increased the concentrations of SO(4), Cl, and Na up to 6000 mg/L and increased the alkalinity up to 923 mg/L CaCO(3) eq. in the coastal aquifer. These waters infiltrated into the tailings deposit at the shelf-tailings interface. Nonremediated tailings had a low-pH oxidation zone (pH 1-4) with significant accumulations of efflorescent salts (10-20 cm thick) at the surface because of upward capillary transport of metal cations in the arid climate. Remediated tailings were characterized by neutral pH and reducing conditions (pH similar to 7, Eh similar to 100 mV). As a result, most bivalent metals such as Cu, Zn, and Ni had very low concentrations (around 0.01 mg/L or below detection limit) because of reduction and sorption processes. In contrast, these reducing conditions increased the mobility of iron from two sources in this system: (1) The originally Fe(III)-rich oxidation zone, where Fe(II) was reduced during the remediation process and formed an Fe(II) plume, and (2) reductive dissolution of Fe(III) oxides present in the original shelf lithology formed an Fe-Mn plume at 10-m depth. These two Fe-rich plumes were pushed toward the shoreline where more oxidizing and higher pH conditions triggered the precipitation of Fe(HI)hydroxide coatings on silicates. These coatings acted as a filter for the arsenic, which naturally infiltrated with the river water (similar to 500 mu g/L As natural background) into the tailings deposit.

Pathways for the synthesis of polyesters in plants; cutin, suberin, and polyhydroxyalkanoates

Plants naturally produce the lipid-derived polyester cutin, which is found in the plant cuticle that is deposited at the outermost extracellular matrix of the epidermis covering nearly all aboveground tissues. Being at the interface between the cell and the external environment, cutin and the cuticle play important roles in the protection of plants from several stresses. A number of enzymes involved in the synthesis of cutin monomers have recently been identified, including several P450s and one acyl-CoA synthetase, thus representing the first steps toward the understanding of polyester formation and, potentially, polyester engineering to improve the tolerance of plants to stresses, such as drought, and for industrial applications. However, numerous processes underlying cutin synthesis, such as a controlled polymerization, still remain elusive. Suberin is a second polyester found in the extracellular matrix, most often synthesized in root tissues and during secondary growth. Similar to cutin, the function of suberin is to seal off the respective tissue to inhibit water loss and contribute to resistance to pathogen attack. Being the main constituent of cork, suberin is a plant polyester that has already been industrially exploited. Genetic engineering may be worth exploring in order to change the polyester properties for either different applications or to increase cork production in other species. Polyhydroxyalkanoates (PHAs) are attractive polyesters of 3-hydroxyacids because of their properties as bioplastics and elastomers. Although PHAs are naturally found in a wide variety of bacteria, biotechnology has aimed at producing these polymers in plants as a source of cheap and renewable biodegradable plastics. Synthesis of PHA containing various monomers has been demonstrated in the cytosol, plastids, and peroxisomes of plants. Several biochemical pathways have been modified in order to achieve this, including the isoprenoid pathway, the fatty acid biosynthetic pathway, and the fatty acid β-oxidation pathway. PHA synthesis has been demonstrated in a number of plants, including monocots and dicots, and up to 40% PHA per gram dry weight has been demonstrated in Arabidopsis thaliana. Despite some successes, production of PHA in crop plants remains a challenging project. PHA synthesis at high level in vegetative tissues, such as leaves, is associated with chlorosis and reduced growth. The challenge for the future is to succeed in synthesis of PHA copolymers with a narrow range of monomer compositions, at levels that do not compromise plant productivity. This goal will undoubtedly require a deeper understanding of plant biochemical pathways and how carbon fluxes through these pathways can be manipulated, areas where plant "omics" can bring very valuable contributions.

Multiple origins of invasive and "native" water frogs (Pelophylax spp.) in Switzerland

The marsh frog (Pelophylax ridibundus) has been introduced in many places of Central and Western Europe due to commercial trades with Eastern Europe, and is rapidly replacing the native pool frog (P. lessonae). A large number of Pelophylax species are distributed in Eastern Europe and the strong phenotypic similarity between these species is rendering their identification hazardous. Consequently, alien populations of Pelophylax might not strictly be composed of P. ridibundus as previously suspected. In the present study, we analyzed the cytochrome b and NADH dehydrogenase subunit 3 genes of introduced and native Pelophylax from Switzerland (299 individuals), in order to properly identify the source populations of the invaders and the genetic status of the native species. Our study highlighted the occurrence of several genetic lineages of invasive frogs in western Switzerland. Unexpectedly, we also showed that several populations of the native pool frog (P. lessonae) cluster with the Italian pool frog P. bergeri from central Italy (considered by some authors as a subspecies of P. lessonae) Hence, these populations are probably also the result of introductions, meaning that the number of native P. lessonae populations is less important than expected in Switzerland. These findings have important implications concerning the conservation of the endemic pool frog populations, as the presence of multiple alien species could strongly affect their long-term subsistence.

SOURCE, FATE AND FUNCTION OF EARLY GLIAL CELLS EXPRESSING NKX2.1 IN MOUSE EMBRYONIC BRAINS

The brain tissue is made of neuronal and glial cells generated in the germinal layer bordering the ventricles. These cells divide, differentiate and migrate following specific pathways. The specification of GABAergic interneurons and glutamatergic neurons has been broadly studied but little is known about the origin, the fate and the function of early glial cells in the embryonic telencephalon. It has been commonly accepted since long that the glial cells and more particularly the astrocytes were generated after neurogenesis from the dorsal telencephalon. However, our work shows that, unlike what was previously thought, numerous glial cells (astroglia and polydendrocytes) are generated during neurogenesis in the early embryonic stages from E14.5 to E16.5, and originate from the ventral Nkx2.1-expressing precursors instead. NK2 homeobox 1 (Nkx2.1) is a member of the NK2 family of homeodomaincontaining transcription factors. The specification of the MGE precursors requires the expression of the Nkx2.1 homeobox gene. Moreover, Nkx2.1 is previously known to regulate the specification of GABAergic interneurons and early oligodendrocytes in the ventral telencephalon. Here, in my thesis work, I have discovered that, in addition, Nkx2.1 also regulates astroglia and polydendrocytes differentiation. The use of Nkx2.1 antibody and Nkx2.1 riboprobe have revealed the presence of numerous Nkx2.1-positive cells that express astroglial markers (like GLAST and GFAP) in the entire embryonic brain. Thus, to selectively fate map MGE-derived GABAergic interneurons and glia, we crossed Nkx2.1-Cre mice, Glast-Cre ERT+/- inducible mice and NG2-Cre mice with the Cre reporter Rosa26-lox-STOP-lox-YFP (Rosa26-YFP) mice. The precise origin of Nkx2.1-positive astroglia has been directly ascertained by combining glial immunostaining and focal electroporation of the pCAG-GS-EGFP plasmids into the subpallial domains of organotypic slices, as well as, by using in vitro neurosphere experiments and in utero electroporation of the pCAG-GS-tomato plasmid into the ventral pallium of E14.5 Nkx2.1-Cre+/Rosa-YFP+/- embryos. We have, thus, confirmed that the three germinal regions of the ventral telencephalon i.e. the MGE, the AEP/POA and the triangular septal nucleus are able to generate early astroglial cells. Moreover, immunohistochemistry for several astroglial cells and polydendrocyte markers, both in the Nkx2.1-/- and control embryos and in the neurospheres, has revealed a severe loss of both glial cell types in the Nkx2.1 mutants. We found that the loss of glia corresponded to a decrease of Nkx2.1-derived precursor division capacity and glial differentiation. There was a drastic decrease of BrdU+ dividing cells labeled for Nkx2.1 in the MGE*, the POA* and the septal nucleus* of Nkx2.1 mutants. In addition, we noticed that while some remaining Nkx2.1+ precursors still succeeded to give rise to post-mitotic neurons in vitro and in vivo in the Nkx2.1-/-, they completely lost the capacity to differentiate in astrocytes. Altogether, these observations indicate for the first time that the transcription factor Nkx2.1 regulates the proliferation and differentiation of precursors in three subpallial domains that generate early embryonic astroglia and polydendrocytes. Furthermore, in order to investigate the potential function of these early Nkx2.1- derived glia, we have performed multiple immunohistochemical stainings on Nkx2.1-/- and wild-type animals, and Nkx2.1-Cre mice that were crossed to Rosa-DTA+/- mice in which the highly toxic diphtheria toxin aided to selectively deplete a majority of the Nkx2.1-derived cells. Interestingly, in these two mutants, we observed a drastic and significant loss of GFAP+, GLAST+, NG2+ and S100ß+ astroglial cells at the telencephalic midline and in the medial cortical areas. This cells loss could be directly correlated with severe axonal guidance defects observed in the corpus callosum (CC), the hippocampal commissure (HIC), the fornix (F) and the anterior commissure (AC). Axonal guidance is a key step allowing neurons to form specific connections and to become organized in a functional network. The contribution of guidepost cells inside the CC and the AC in mediating the growth of commissural axons have until now been attributed to specialized midline guidepost astroglia. Previous published results in our group have unravelled that, during embryonic development, the CC is populated in addition to astroglia by numerous glutamatergic and GABAergic guidepost neurons that are essential for the correct midline crossing of callosal axons. Therefore, the relative contribution of individual neuronal or glial populations towards the guidance of commissural axons remains largely to be investigated to understand guidance mechanisms further. Thus, we crossed Nkx2.1-Cre mice with NSE-DTA+/- mice that express the diphtheria toxin only in neurons and allowed us to selectively deplete Nkx2.1-derived GABAergic neurons. Interestingly, in the Nkx2.1-/- mice, the CC midline was totally disorganized and the callosal axons partly lost their orientation, whereas in the Nkx2.1Cre+/Rosa-DTA+/- and the Nkx2.1Cre+/NSE-DTA+/- mice, the axonal organization of the CC was not affected. In the three types of mice, hippocampal axons of the fornix were not properly fasciculated and formed disoriented bundles through the septum. Additionally, the AC formation was completely absent in Nkx2.1-/- mice and the AC was divided into two/three separate paths in the Nkx2.1Cre+/Rosa-DTA+/- mice that project in wrong territories. On the other hand, the AC didn't form or was reduced to a relatively narrower tract in the Nkx2.1Cre+/NSE-DTA+/- mice as compared to wild-type AC. These results clearly indicate that midline Nkx2.1-derived cells play a major role in commissural axons pathfinding and that both Nkx2.1-derived guidepost neurons and glia are necessary elements for the correct development of these commissures. Furthermore, during our investigations on Nkx2.1-/- and Nkx2.1Cre+/Rosa-DTA+/- mice, we noticed similar and severe defects in the erythrocytes distribution and the blood vessels network morphology in the embryonic brain of both mutants. As the Cre-mediated recombination was never observed to occur in the blood vessels of Nkx2.1-Cre mice, we inferred that the vessels defects observed were due to the loss of Nkx2.1-derived cells and not to the cells autonomous effects of Nkx2.1 in regulating endothelial cell precursors. Thereafter, the respective contribution of individual Nkx2.1-regulated neuronal or glial populations in the blood vessels network building were studied with the use of transgenic mice strains. Indeed, the use of Nkx2.1Cre+/NSE-DTA+/- mice indicated that the Nkx2.1-derived neurons were not implicated in this process. Finally, to discriminate between the two Nkx2.1-derived glial cell populations, the GLAST+ astroglia and the NG2+ polydendrocytes, an NG2-Cre mouse strain crossed to the Rosa-DTA+/- mice was used. In that mutant, the blood vessel network and the erythrocytes distribution were similarly affected as observed in Nkx2.1Cre+/Rosa-DTA+/- animals. Therefore, this result indicates that most probably, the NG2+ polydendrocytes are involved in helping to build the vessels network in the brain. Taken altogether, these observations show that during brain development, Nkx2.1- derived embryonic glial cells act as guidepost cells on the guidance of axons as well as forming vessels. Both Nkx2.1-regulated guidepost GABAergic neurons and glia collaborate to guide growing commissural axons, while polydendrocytes are implicated in regulating brain angiogenesis. - Le tissu cérébral est composé de cellules neuronales et gliales générées dans les couches germinales qui bordent les ventricules. Ces cellules se divisent, se différencient et migrent selon des voies particulières. La spécification des interneurones GABAergiques et des neurones glutamatergiques a été largement étudiée, par contre, l'origine, le destin et la fonction des cellules gliales précoces du télencéphale embryonnaire restent peu élucidées. Depuis longtemps, il était communément accepté que les cellules gliales, et plus particulièrement les astrocytes, sont générés après la neurogénèse à partir du télencéphale dorsal. Toutefois, notre travail montre que de nombreuses cellules gliales sont générées à partir de précurseurs ventraux qui expriment le gène Nkx2.1, entre E14.5 et E16.5, c'est-à dire,à des stades embryonnaires très précoces. Le gène NK2 homéobox 1 (Nkx2.1) appartient à une famille de facteurs de transcription appelée NK2. Il s'agit de protéines qui contiennent un homéo-domaine. La spécification des précurseurs de la MGE requiert l'expression du gène homéobox Nkx2.1. De plus, la fonction du gène Nkx2.1 dans la régulation de la spécification des interneurones GABAergiques et des oligodendrocytes dans le télencéphale ventral était déjà connue. Au cours de mon travail de thèse, j'ai également mis en évidence que, Nkx2.1 régule aussi les étapes de prolifération et de différenciation de divers sous-types de cellules gliales soit de type astrocytes ou bien polydendrocytes. L'utilisation d'un anticorps contre la protéine Nkx2.1 ainsi qu'une sonde à ribonucléotides contre l'ARN messager du gène Nkx2.1 ont révélé la présence de nombreuses cellules positives pour Nkx2.1 qui exprimaient des marqueurs astrocytaires (comme GLAST et GFAP) dans le télencéphale embryonnaire. Afin de déterminer de manière sélective le sort des interneurones GABAergiques, des polydendrocytes et des astrocytes dérivés de la MGE, nous avons croisé soit des souris Nkx2.1-Cre, des souris Glast-Cre ERT+/- inductibles ou bien des souris NG2-Cre avec des souris Rosa26-lox-STOP-lox-YFP (Rosa26-YFP) Cre rapportrices. L'origine précise des astroglies positives pour Nkx2.1 a été directement établie en combinant une coloration immunologique pour les glies et une électroporation focale d'un plasmide pCAG-GS-EGFP dans les domaines subpalliaux de tranches organotypiques, puis également, par des cultures de neurosphères in vitro et des expériences d'électroporation in utero d'un plasmide pCAG-GS-tomato dans le pallium ventral d'embryons Nkx2.1-Cre+/Rosa- YFP+/- au stade E14.5. Nous avons donc confirmé que les trois régions germinales du télencéphale ventral, c'est-à-dire, la MGE, l'AEP/POA et le noyau triangulaire septal sont capables de générer des cellules astrogliales. D'autre part, l'immunohistochimie pour plusieurs marqueurs d'astrocytes ou de polydendrocytes, dans les embryons Nkx2.1-/- et contrôles ainsi que dans les neurosphères, a révélé une sévère perte de ces deux types gliaux chez les mutants. Nous avons trouvé que la perte de glies correspondait à une diminution de la capacité de division des précurseurs dérivés de Nkx2.1, ainsi que l'incapacité de ces précurseurs de se différencier en cellules gliales. Nous avons en effet observé une diminution importante des cellules BrdU+ en division exprimant Nkx2.1dans la MGE*, la POA* et le noyau septal* des mutants pour Nkx2.1. D'autre part, nous avons pu mettre en évidence aussi bien in vitro, qu'in vivo, que certains précurseurs Nkx2.1+ chez le mutant gardent la capacité à se différencier en neurones tandis qu'ils perdent celle de se différencier en cellules gliales. Prises dans leur ensemble, ces observations indiquent pour la première fois que le facteur de transcription Nkx2.1 régule les étapes de prolifération et de différentiation des précurseurs des trois domaines subpalliaux qui génèrent les astroglies et polydendrocytes embryonnaires précoces. Par la suite, dans le but de comprendre la fonction potentielle de ces glies précoces, nous avons procédé à de multiples colorations immunohistochimiques sur des animaux Nkx2.1-/- et sauvages, ainsi que sur des souris Nkx2.1-Cre croisées à des souris Rosa-DTA+/- dans lesquelles la toxine diphthérique hautement toxique a permis de supprimer sélectivement la majorité des cellules dérivées de Nkx2.1. De manière intéressante, nous avons observé dans ces deux mutants, une perte drastique et significative de cellules astrogliales GFAP+, GLAST+ et polydendrocytaires NG2+ et S100ß+ dans le télencéphale, à la midline et dans les aires corticales médianes. Ces pertes ont pu être directement corrélées avec des défauts de guidage axonal observés dans le corps calleux (CC), la commissure hippocampique (HIC), le fornix (F) et la commissure antérieure (AC). Le guidage axonal est une étape clé permettant aux neurones de former des connections spécifiques et de s'organiser dans un réseau fonctionnel. La contribution des cellules « guidepost » dans le CC et dans la AC comme médiateurs de la croissance des axones commissuraux à jusqu'à aujourd'hui été attribuée spécifiquement à des astroglies « guidepost » de la midline. Des résultats publiés précédemment dans notre groupe, ont permis de montrer que, pendant le développement embryonnaire, le CC est peuplé en plus de la glie par de nombreux neurones « guidepost » glutamatergiques et GABAergiques qui sont essentiels pour le croisement correct des axones callosaux à la midline. Ainsi, la contribution relative des populations individuelles neuronales ou gliales pour le guidage des axones commissuraux demande à être approfondie afin de mieux comprendre les mécanismes de guidage. A ces fins, nous avons croisé des souris Nkx2.1-Cre avec des souris NSE-DTA+/- qui expriment la toxine diphthérique uniquement dans les neurones et ainsi, nous avons pu sélectivement supprimer les neurones dérivés de domaines Nkx2.1+. Dans les souris Nkx2.1-/-,nous avons découvert que le CC était désorganisé avec des axones callosaux perdant partiellement leur orientation, alors que dans les souris Nkx2.1Cre+/Rosa-DTA+/- et Nkx2.1Cre+/NSE-DTA+/-, l'organisation axonale n'était pas affectée. De plus, les faisceaux hippocampiques du fornix étaient défasciculés dans les trois types de mutants. Par ailleurs, la formation de la commissure antérieure (AC) était complètement absente dans les souris Nkx2.1-/- d'une part, et d'autre part, celle-ci était divisée en deux à trois voies séparées dans les souris Nkx2.1Cre+/Rosa-DTA+/-. Finalement, la AC était soit absente, soit réduite de manière ne former plus qu'un faisceau relativement plus étroit dans les souris Nkx2.1Cre+/NSE-DTA+/- en comparaison avec la AC sauvage. Ces derniers résultats indiquent clairement que les cellules dérivées de Nkx2.1 à la midline, jouent un rôle majeur dans le guidage des axones commissuraux et que, autant les neurones, que les astrocytes « guidepost » dérivés de Nkx2.1, sont des éléments nécessaires au développement correct de ces commissures. En outre, lors de nos investigations sur les souris Nkx2.1-/- et Nkx2.1Cre+/Rosa-DTA+/-, nous avons remarqués des défauts sévères et similaires dans la distribution des erythrocytes et dans la morphologie du réseau de vaisseaux sanguins dans le cerveau embryonnaire des deux mutants précités. Puisque nous n'avons jamais observé de recombinaison de la Cre recombinase dans les vaisseaux sanguins des souris Nkx2.1Cre, nous en avons déduit que les défauts de vaisseaux observés étaient dus à la perte de cellules dérivées de Nkx2.1. Il existerait donc en plus de la fonction cellulaire autonome de Nkx2.1 reconnue pour régulée directement la spécification des cellules endothéliales, une fonction indirecte de Nkx2.1. Afin de déterminer la contribution respective des populations individuelles neuronales ou gliales régulées par Nkx2.1 dans la construction du réseau de vaisseaux sanguins, nous avons utilisé diverses lignées de souris transgéniques. L'utilisation de souris Nkx2.1Cre+/NSE-DTA+/- a indiqué que les neurones dérivés de Nkx2.1 n'étaient pas impliqués dans ce processus. Finalement, afin de discriminer entre les deux populations de cellules gliales dérivées de Nkx2.1, les astroglies et les polydendrocytes, nous avons croisé une lignée de souris NG2-Cre avec des souris Rosa-DTA+/-. Dans ce dernier mutant, le réseau de vaisseaux sanguins du cortex ainsi que la distribution des erythrocytes étaient affectés de la même manière que dans le cortex des souris Nkx2.1Cre+/Rosa-DTA+/-. Par conséquent, ce résultat indique que très probablement, les polydendrocytes NG2+ sont impliqués dans la mise en place du réseau de vaisseaux dans le cerveau. Prises dans leur ensemble, ces observations montrent que durant le développement embryonnaire du cerveau, des sous-populations de glies régulées par Nkx2.1 jouent un rôle de cellules « guidepost » dans le guidage des axones, ainsi que des vaisseaux. Les polydendrocytes sont impliquées dans la régulation de l'angiogenèse tandis que, autant les neurones GABAergiques que les astrocytes collaborent dans le guidage des axones commissuraux en croissance.