Radiometrically determined dates and sedimentation rates for recent sediments in nine North African wetland lakes (the CASSARINA Project)

Sediment cores were collected from nine wetland lakes in Morocco, Tunisia and Egypt for the CASSARINA project investigating environmental change in Northern African wetlands. The cores were dated radiometrically by using natural (²¹⁰Pb) and artificial (¹³⁷Cs and ²⁴¹Am) radionuclides. At sites in Morocco and Tunisia with mean annual rainfall totals ranging from 500–1000 mm yr^–1, fallout records were generally satisfactory and it was possible to develop independent sediment chronologies based on the radiometric data alone. At the Egyptian sites, rainfall was less than 200 mm yr^–1 and fallout records were much less distinct. At these sites the radiometric data could only be used to give an indication of mean sedimentation rates during the past 30–40 years. By using a combination of fallout radionuclide, pollen, and macrofossil stratigraphic records it was however possible to determine a credible sediment chronology spanning the major part of the 20th century. Applying this chronology to records of spheroidal carbonaceous particles (SCP) from the same sediment cores, the onset of significant levels of atmospheric pollution in the Nile Delta is dated in all three cores to the mid 1950s. Results from a number of lakes (Sidi Bou Rhaba, Ichkeul and Korba) revealed high and accelerating siltation rates, threatening their continued existence beyond the next few decades. In contrast, sedimentation rates at all three Nile Delta sites appear to have declined in recent decades, most probably due to the impact of the Nile barrages.     

Biochemical and Immunocytological Characterizations of Arabidopsis Pollen Tube Cell Wall

During plant sexual reproduction, pollen germination and tube growth require development under tight spatial and temporal control for the proper delivery of the sperm cells to the ovules. Pollen tubes are fast growing tip-polarized cells able to perceive multiple guiding signals emitted by the female organ. Adhesion of pollen tubes via cell wall molecules may be part of the battery of signals. In order to study these processes, we investigated the cell wall characteristics of in vitro-grown Arabidopsis (Arabidopsis thaliana) pollen tubes using a combination of immunocytochemical and biochemical techniques. Results showed a well-defined localization of cell wall epitopes. Low esterified homogalacturonan epitopes were found mostly in the pollen tube wall back from the tip. Xyloglucan and arabinan from rhamnogalacturonan I epitopes were detected along the entire tube within the two wall layers and the outer wall layer, respectively. In contrast, highly esterified homogalacturonan and arabinogalactan protein epitopes were found associated predominantly with the tip region. Chemical analysis of the pollen tube cell wall revealed an important content of arabinosyl residues (43%) originating mostly from (1→5)-α-l-arabinan, the side chains of rhamnogalacturonan I. Finally, matrix-assisted laser desorption ionization time-of-flight mass spectrometry analysis of endo-glucanase-sensitive xyloglucan showed mass spectra with two dominant oligosaccharides (XLXG/XXLG and XXFG), both being mono O-acetylated, and accounting for over 68% of the total ion signals. These findings demonstrate that the Arabidopsis pollen tube wall has its own characteristics compared with other cell types in the Arabidopsis sporophyte. These structural features are discussed in terms of pollen tube cell wall biosynthesis and growth dynamics.Fertilization of flowering plants requires the delivery of the two sperm cells, carried by a fast growing tip-polarized pollen tube, to the egg cell. In plants with dry stigma and solid style such as Arabidopsis (Arabidopsis thaliana), this process begins with the deposition and specific adhesion of the pollen grains on the stigmatic tissue, subsequent hydration of the pollen grains, and germination of pollen tubes (Palanivelu and Preuss, 2000). Pollen tubes invade the papillae cell wall of the stigma, enter the short style, and grow through the apoplast of the specialized transmitting tract (TT) that is filled with a nutrient-rich extracellular matrix (Kandasamy et al., 1994; Lennon et al., 1998). During this invasive growth, pollen tubes are guided to the ovules via signals that need to pass through the cell wall to reach their membrane-associated or intracellular targets (Lord and Russell, 2002; Kim et al., 2003; Boavida et al., 2005; McCormick and Yang, 2005; Johnson and Lord, 2006). In plant species with wet stigma and hollow style such as lily (Lilium longiflorum), adhesion between the pollen tube wall and the TT epidermis extracellular matrix is important for the growth of the pollen tubes toward the ovules (Mollet et al., 2000, 2007; Park et al., 2000; Chae et al., 2007). In addition to being the interface between the tube cells and the surroundings (female sporophyte or culture medium), the pollen tube wall also controls the cell shape, protects the generative cells, and allows resistance against turgor pressure (Geitmann and Steer, 2006; Geitmann, 2010).Most of our knowledge on cell wall polymers of higher plants comes from investigations on vegetative organs in which cells have diffuse growth. The cell wall is mainly composed of polysaccharides (cellulose, hemicellulose, pectin, and occasionally callose, depending on the tissue) and proteoglycans (e.g. extensin and arabinogalactan proteins [AGPs]) forming a complex network with processing enzymes.Pectins are complex wall macromolecules with uncertain supramolecular organization (Vincken et al., 2003) consisting of homogalacturonan (HG) that can be methylesterified and acetylesterified, rhamnogalacturonan I (RG-I), rhamnogalacturonan II (RG-II), and xylogalacturonan (Carpita and McCann, 2000). HG is a polymer of repeated units of (1→4)-α-d-GalUA that can be cross-linked with calcium upon block-wise action of pectin methylesterases (PMEs) on methylesterified HG (Micheli, 2001). RG-II has the same homopolymer backbone as HG but is substituted with four different oligosaccharides composed of unusual sugars, such as apiose, aceric acid, and 3-deoxy-d-manno-2-octulosonic acid, of unknown function (for review, see Caffall and Mohnen, 2009). RG-I consists of the repeating disaccharide (1→4)-α-d-GalUA-(1→2)-α-l-Rha, with a wide variety of side chains attached to the rhamnosyl residues, ranging from monomers to large oligosaccharides such as (1→4)-β-d-galactan, (1→5)-α-l-arabinan, and/or type I arabinogalactan (Caffall and Mohnen, 2009).Xyloglucan (XyG) is the major hemicellulosic polysaccharide of the primary wall of flowering plants. Classic XyG consists of a (1→4)-β-d-glucan backbone substituted with Xyl, Gal-Xyl, or Fuc-Gal-Xyl motifs, which correspond, according to the one-letter code proposed by Fry et al. (1993), to X, L, and F, respectively, G being the unsubstituted glucosyl residue of the glucan backbone. The main XyG fragments released after endo-glucanase treatment of the cell wall from wild-type Arabidopsis vegetative organs are generally XXXG, XXLG/XLXG, XXFG, and XLFG (Zablackis et al., 1995; Lerouxel et al., 2002; Nguema-Ona et al., 2006; Obel et al., 2009). In addition, O-acetylation of XyG can occur, most generally on the galactosyl residues, but its biological function is unknown (Cavalier et al., 2008). In the primary wall, XyG interacts with cellulose microfibrils via hydrogen bonds and participates in the control of cell expansion (Cosgrove, 1999).AGPs and extensin belong to the Hyp-rich glycoproteins superfamily with very high levels of type II arabinogalactan glycosylation (Nothnagel, 1997; Showalter, 2001). These proteoglycans have been implicated in many aspects of plant development, including cell expansion, cell signaling and communication, embryogenesis, wound response, and pollen tube guidance (Wu et al., 1995; Nothnagel, 1997; Seifert and Roberts, 2007; Driouich and Baskin, 2008).Despite the importance of pollen tubes for the delivery of the sperm cells to the egg, little is known about the underlying molecular mechanisms that regulate the mechanical interaction of pollen tubes with female floral tissues. There are very scarce data concerning the different components of the pollen tube cell wall. Past approaches to characterize the pollen tube cell wall are limited to a few plant genera, including Camellia (Nakamura and Suzuki, 1981), Lilium (Jauh and Lord, 1996; Mollet et al., 2002), Nicotiana (Rae et al.,1985; Li et al., 1995; Ferguson et al., 1998; Qin et al., 2007), Pinus (Derksen et al., 1999), and Zea (Rubinstein et al., 1995), and are mostly based on immunocytochemistry. These studies revealed that, depending on the species, the pollen tube cell wall contains epitopes that are found in the polymers described above, including HGs with varying levels of methylesterification, AGPs, extensin-like proteins, and low amounts of cellulose. Unlike most other plant cells, callose, a (1→3)-β-glucan, is predominant and is deposited in the wall back from the tip. Moreover, it is deposited at regular intervals to form callose plugs that maintain the tube cell in the apical expanding region of the tube and separate the viable from the degenerating region of the tube (for review, see Geitmann and Steer, 2006). Only a few reports have investigated the pollen tube of the model plant Arabidopsis. They have focused either on in vivo-grown or on in vitro-grown pollen tubes using monoclonal antibodies (MAbs) directed against a subset of cell wall epitopes present in HG, XyG, and AGPs (Lennon and Lord, 2000; Freshour et al., 2003; Pereira et al., 2006), but quantitative chemical analyses are lacking. This lack of information is most likely due to the fact that substantial amounts of pollen tube material are needed for chemical analysis, and a reproducible and efficient method for liquid culture of Arabidopsis pollen tubes had not been established until recently (Boavida and McCormick, 2007; Bou Daher et al., 2009).Here, we report the composition and localization of different cell wall polymers of in vitro-grown wild-type Arabidopsis pollen tubes based on biochemical analyses coupled to immunocytochemical investigations both at light and transmission electron microscopy (TEM) levels using recently developed MAbs. Our results show distinct patterns of labeling (tip, whole tube, and shank of the tube) depending on the recognized epitope. The most striking observations are (1) the abundance of (1→5)-α-l-arabinan in the tube wall (greater than 40 mol % of Ara), mostly localized, with LM6 and LM13, in the outer wall layer of the tube and (2) an atypical XyG matrix-assisted laser desorption ionization time-of-flight mass spectrometry (MALDI-TOF MS) profile with over 68% of the oligosaccharide fragments being O-acetylated.     

Open water zooplankton communities in North African wetland lakes: the CASSARINA Project

Zooplankton (Copepoda, Cladocera, Ostracoda, Rotifera and Diptera larvae) in nine North African lakes was collected from open water areas over twenty months during 1997/99. The results were used to monitor changes in the pelagic micro-invertebrate fauna of these sites with the purpose of exploring diversity structure and regional species occurrences.The studied sites formed three distinct groups based on hydrology and water quality criteria: (i) acid water with no marine connection (Megene Chitane); (ii) alkaline freshwater/brackish with no marine connection (Merja Sidi Bou Rhaba and Merja Bokka); (iii) freshwater/brackish with marine connection (Merja Zerga, Lac de Korba, Garaet El Ichkeul and three Nile Delta lakes). However, cluster analysis of the zooplankton data alone indicated four groups with Korba being separated because of its prevalence of species tolerant of summer hypersalinity.The total regional zooplanktonic species richness found was 88 taxa and these were characterized by species tolerant of widely fluctuating environmental conditions. However, some recorded species were very rare for North African freshwaters (e.g. Alonella excisa, Leydigia quadrangularis and, Ilyocryptus sordidus) and generally indicate favourable environmental conditions of low salinity and temperature. The sites influenced by marine waters generally exhibited slightly lower numbers of species but which generally demonstrate cosmopolitan distributions. Distinct seasonal patterns in species distributions were more similar to those observed in European lakes rather than to those of lower latitudes sites.Zooplankton play a key role in maintaining aquatic ecosystem quality in the North African study lakes and the community distributions described for the late 20th century help set biodiversity base-line data for future studies. If the remaining wetland lakes in this region are to persist as important resources during the 21st century, they will need to be managed in a way that ensures that aquatic diversity is maintained.     

Application of remote sensing to site characterisation and environmental change analysis of North African coastal lagoons

This article describes the use of satellite imagery for identifying key environmental characteristics within three North African coastal lagoons (Merja Zerga, Morocco; Ghar El Melh, Tunisia and Lake Manzala, Egypt) and for detecting the major environmental changes within these environments. A combination of Landsat MSS, Landsat TM, Landsat ETM+ and ASTER imagery was acquired for the three sites for a period covering the last three decades (1972–2004). Following geometric correction and enhancement, the interpretation of the most recent image acquired for each of the three lagoons provides important insights into their current conditions. For Merja Zerga, these include the distribution of the largest channels which drain extensive inter-tidal mudflats and the two major depositional features associated with sources of freshwater. The distribution of marginal aquatic vegetation is highlighted as is the intensive use of the surrounding landscape for agriculture. Intensive agriculture around Ghar El Melh is also indicated. The influence of the Mejerda River, which was diverted away from the lagoon over 100 years ago, is shown to persist as a residual area of deltaic deposits in shallow water that has been eroded over time. Coastal processes including the direction of the alongshore sediment transport and the influence of engineering work associated with port construction can also be recognised. Within Lake Manzala, vegetated islands divide the lake into a series of sub-basins which can be clearly distinguished. The large influence of human activities within this lake can be identified and include reclamation for agriculture and the conversion of parts of the lake bed for fish farms. The historical images available for the three lagoons provide important insights into decadal scale changes, which have been greatest at Lake Manzala. Since the early 1970s large parts of the lake, in particular in the southwest where the shoreline has migrated northwards, have been reclaimed. Major engineering works, such as the El Salam Canal and road embankments, are shown to have resulted in significant lake change. The distribution of emergent vegetation within the lake has also changed. Classification of images for this lake into open water, vegetation and land enables the quantification of these changes. Between 1973 and 2003, the lake declined in area by approximately 50%. Changes at Merja Zerga over the last three decades include reconfiguration of the marine outlet and the expansion of the internal delta at the end of the Nador Canal. The images of this site clearly demonstrate the intensification of agriculture around the lagoon. The most marked changes evident within the images of Ghar El Melh concern the sand bars that separate the lagoon from the sea. Geomorphological processes operating within the coastal zone have resulted in the straightening of the bars with central sections migrating out towards the sea. Remote sensing is established as a promising application for detecting the quantitative surface cover changes in coastal lagoons and their near landscapes. Guest editors: J. R. Thompson & R. J. Flower Hydro-ecological Monitoring and Modelling of North African Coastal Lagoons     

Recent environmental change in North African wetland lakes: a baseline study of organochlorine contaminant residues in sediments from nine sites in the CASSARINA Project

Sediment from 9 lakes and lagoons in 3 North African countries was analysed for a suite of organochlorine insecticides and polychlorinated biphenyls (PCBs). Residues of -HCH and pp-DDE were detected in most cores, and radiometric dating of the sediment enabled time profiles to be constructed which are indicative of recent use of the insecticide lindane and previous use of the insecticide DDT in the region. Absolute concentrations of these pesticides were relatively low compared to reported values for other world-wide locations, but exceeded recommended Canadian and Dutch environmental quality standards at several sites. Maximum fluxes of -HCH and pp-DDE (190 and 95 g m^–2 yr^–1, respectively) were relatively high and comparable to some sediments in North America and the United Kingdom. Other organochlorine pesticides including dieldrin were detected at low levels in some samples. Selected PCB congeners were detected at trace levels at 2 sites only, indicating low levels of industrial contamination at the sampling locations. Tropical and sub-tropical regions of the world have been suggested to be contemporary sources of globally distributed organochlorine contaminants. These data are discussed with respect to this hypothesis.     

Subcompartment localization of the side chain xyloglucan-synthesizing enzymes within Golgi stacks of tobacco suspension-cultured cells

Xyloglucan is the dominant hemicellulosic polysaccharide of the primary cell wall of dicotyledonous plants that plays a key role in plant development. It is well established that xyloglucan is assembled within Golgi stacks and transported in Golgi-derived vesicles to the cell wall. It is also known that the biosynthesis of xyloglucan requires the action of glycosyltransferases including α-1,6-xylosyltransferase, β-1,2-galactosyltransferase and α-1,2-fucosyltransferase activities responsible for the addition of xylose, galactose and fucose residues to the side chains. There is, however, a lack of knowledge on how these enzymes are distributed within subcompartments of Golgi stacks. We have undertaken a study aiming at mapping these glycosyltransferases within Golgi stacks using immunogold-electron microscopy. To this end, we generated transgenic lines of tobacco (Nicotiana tabacum) BY-2 suspension-cultured cells expressing either the α-1,6-xylosyltransferase, AtXT1, the β-1,2-galactosyltransferase, AtMUR3, or the α-1,2-fucosyltransferase AtFUT1 of Arabidopsis thaliana fused to green-fluorescent protein (GFP). Localization of the fusion proteins within the endomembrane system was assessed using confocal microscopy. Additionally, tobacco cells were high pressure-frozen/freeze-substituted and subjected to quantitative immunogold labelling using anti-GFP antibodies to determine the localization patterns of the enzymes within subtypes of Golgi cisternae. The data demonstrate that: (i) all fusion proteins, AtXT1-GFP, AtMUR3-GFP and AtFUT1-GFP are specifically targeted to the Golgi apparatus; and (ii) AtXT1-GFP is mainly located in the cis and medial cisternae, AtMUR3-GFP is predominantly associated with medial cisternae and AtFUT1-GFP mostly detected over trans cisternae suggesting that initiation of xyloglucan side chains occurs in early Golgi compartments in tobacco cells.     

North African wetland lakes: characterization of nine sites included in the CASSARINA Project

Exploitation of land and water resources has increased rapidly in North Africa during the 20th century, paralleling regional population growth. As part of the CASSARINA Project (see Flower, 2001), the environmental status of nine wetland lakes in Morocco, Tunisia, and Egypt was evaluated. All are conservationally important habitats and several are Ramsar Sites (internationally recognized bird reserves) and several support significant fisheries. All are shallow (<2 m in depth) but vary greatly in area.Where available, documentary information on relevant 20th century changes is given. Survey transects for aquatic vegetation were established and used to provide baseline ecological information on the aquatic plant communities during 1997–1999. Unusually, one site (Tunisian Megene Chitane) supported acidophilous vegetation (some taxa being nationally rare). Aquatic macrophytes declined catastrophically at two sites during the 1990s. Merja Bokka was drained in 1998 and, at Garaet El Ichkeul, fringing Phragmites and Scirpus spp. were lost, mainly as a result of salinity changes. Elsewhere, fringing macrophytes remain (extensively so in the Nile Delta lakes) common, despite major land reclamation and water quality problems, or are degraded by grazing (Merja Zerga). Marginal vegetation during 1997/98 changed markedly at Megene Chitane due to water level lowering.Documentary records indicated that throughout the 20th century, reclamation and hydrologic modifications, mainly for agricultural purposes, affected all nine sites. The loss of lake area by reclamation is substantial for the Nile Delta lakes (Edku, Burullus and Manzala). For the western sites, some data indicate increasing salinity in the most recent decade but the Delta lakes have become generally fresher during the 20th century, as supply of Nile water for irrigation increased.Despite intense human disturbance, many of the remaining CASSARINA sites still support regions of high aquatic diversity. Spatial scale monitoring of the larger sites for seasonal and inter-annual changes in open water area and in aquatic plant abundances is a key requirement for integrated environmental change assessment in the 21st century.     

Both mating types of the wheat pathogen Mycosphaerella graminicola are present in Morocco

Septoria tritici blotch caused by the heterothallic ascomycete Mycosphaerella graminicola is one of the most currently damaging diseases on wheat crops worldwide. So far, no information was reported about the status of sexual reproduction of this pathogen under Moroccan conditions. We investigated here for the first time the occurrence of the two mating types (MAT1-1 and MAT1-2) of M. graminicola in Morocco by sampling 141 single-conidial isolates from 4 important wheat producing regions (Gharb, Sa?s, Chaouia and Tadla). The mating type of each isolate was determined by amplification with multiplex PCR of a partial sequence from the corresponding idiomorph. Overall, 43% out of the assessed isolates were MAT1-1 and 57 % were MAT1-2. Both mating types were identified within the 3 sampled regions Gharb, Sa?s and Chaouia, but not in Tadla, where only MAT1-2 isolates were found. The presence of the two mating types highlighted here offers a suitable genetic condition for M. graminicola to occur sexual reproduction in Morocco. The potential of sexual recombination will be examined by the study of mating type frequencies using a large sample size as well as by searching and quantification of pseudothecia in the field.     

Hydrological characteristics of three North African coastal lagoons: insights from the MELMARINA project

Hydrological and hydrodynamic characteristics are important controls in all wetlands including coastal lagoons. Enhanced understanding of lagoon functioning can be obtained through the acquisition and interpretation of hydrological, meteorological and related data. The MELMARINA Project investigated links between hydrological and ecological conditions within North African coastal lagoons. It employed three primary sites: Merja Zerga in Morocco, Ghar El Melh in Tunisia and Lake Manzala in Egypt. Hydrological, meteorological and related data were acquired for these lagoons. Data included precipitation, evaporation, wind speed and direction, freshwater discharges into the lagoons, tides beyond the lagoons in the open sea, lagoon bathymetry and time series of lagoon water levels/depths. Data were acquired from secondary sources (including online archives) and targeted field survey and monitoring programmes. Interpretation of these data provides insights into the hydrological functioning of the lagoons and contributed to the modelling requirements of MELMARINA. The functioning of Merja Zerga is dominated by the exchange of water between the Atlantic and the lagoon. Large, tidally induced oscillations in water level are responsible for the inundation of extensive inter-tidal mudflats whilst the rapid replacement of water by exchanges with the sea diminishes the influence of freshwater inflows and winter rainfall. The smaller Mediterranean tides reduce the magnitude of lagoon–sea exchanges and result in much smaller water level oscillations within Ghar El Melh. As a result, this lagoon lacks the inter-tidal environments found within the Moroccan site and hydrodynamic conditions are more likely influenced by wind set up. Although freshwater inflows to Ghar El Melh are smaller than those in Merja Zerga, their effects, coupled with heavy winter rainfall, can persist due to longer residence times of water within the lagoon. Freshwater inflows are central to the functioning of Lake Manzala. Large discharges of relatively freshwater are provided by major drains which cross the Nile Delta. These influence lake water levels and are responsible for the freshwater conditions within large parts of the lake. The small tidal range of the eastern Mediterranean, coupled with constricted connections with the sea, reduces the influence of lake–sea exchanges and tidally induced water level oscillations within Lake Manzala. Guest editors: J. R. Thompson & R. J. Flower Hydro-ecological Monitoring and Modelling of North African Coastal Lagoons