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Dew formation was investigated in three developmental stages of biological soil crusts (BSC), which were collected along a catena of an inland dune and in the initial substrate. The Penman equation, which was developed for saturated surfaces, was modified for unsaturated surfaces and used for prediction of dewfall rates. The levels of surface saturation required for this approach were predicted using the water retention functions and the thicknesses of the BSCs. During a first field campaign (2–3 August 2011), dewfall increased from 0.042 kg m−2 for the initial sandy substrate to 0.058, 0.143 and 0.178 kg m−2 for crusts 1 to 3, respectively. During a second field campaign (17–18 August 2011), where dew formation was recorded in 1.5 to 2.75-h intervals after installation at 21:30 CEST, dewfall increased from 0.011 kg m−2 for the initial sandy substrate to 0.013, 0.028 and 0.055 kg m−2 for crusts 1 to 3, respectively. Dewfall rates remained on low levels for the substrate and for crust 1, and decreased overnight for crusts 2 and 3 (with crust 3 > crust 2 > crust 1 throughout the campaign). Dew formation was well reflected by the model response. The suggested mechanism of dew formation involves a delay in water saturation in near-surface soil pores and extracellular polymeric substances (EPS) where the crusts were thicker and where the water capacity was high, resulting in elevated vapor flux towards the surface. The results also indicate that the amount of dewfall was too low to saturate the BSCs and to observe water flow into deeper soil. Analysis of the soil water retention curves revealed that, despite the sandy mineral matrix, moist crusts clogged by swollen EPS pores exhibited a clay-like behavior. It is hypothesized that BSCs gain double benefit from suppressing their competitors by runoff generation and from improving their water supply by dew collection. Despite higher amounts of dew, the water availability to the crust community decreases with crust development, which may be compensated by ecophysiological adaptation of crust organisms, and which may further suppress higher vegetation or mosses.
Succession of N Cycling Processes in Biological Soil Crusts on a Central European Inland Dune
(2013)
Biological soil crusts (BSCs) are microbial assemblages that occur worldwide and facilitate ecosystem development by nitrogen (N) and carbon accumulation. N turnover within BSC ecosystems has been intensively studied in the past; however, shifts in the N cycle during BSC development have not been previously investigated. Our aim was to characterise N cycle development first by the abundance of the corresponding functional genes (in brackets) and second by potential enzyme activities; we focussed on the four processes: N fixation (nifH), mineralisation as proteolysis and chitinolysis (chiA), nitrification (amoA) and denitrification (nosZ). We sampled from four phases of BSC development and from a reference located in the rooting zone of Corynephorus canescens, on an inland dune in Germany. BSC development was associated with increasing amounts of chlorophyll, organic carbon and N. Potential activities increased and were highest in developed BSCs. Similarly, the abundance of functional genes increased. We propose and discuss three stages of N process succession. First, the heterotrophic stage (mobile sand without BSCs) is dominated by mineralisation activity. Second, during the transition stage (initial BSCs), N accumulates, and potential nitrification and denitrification activity increases. Third, the developed stage (established BSCs and reference) is characterised by the dominance of nitrification.
In hot and wet conditions in summer, the biological soil crusts (BSCs) decreased soil temperature by up to 11.8 °C, 7.5 °C, 5.4 °C, and 3.2 °C at surface, 5 cm, 15 cm, and 30 cm, respectively; while in cold and dry conditions in winter the BSCs increased soil temperature by up to 1.2 °C, 1.2 °C, and 1.1 °C at 5 cm, 15 cm, and 30 cm, respectively. The daily mean soil temperatures of the BSCs in a whole year were averagely increased by 0.57 ± 0.04 °C, 0.31 ± 0.04 °C, and 0.22 ± 0.04 °C at 5 cm, 15 cm, and 30 cm, respectively. The effects of the BSCs on soil temperature were positively correlated with air temperature and soil moisture, and decreased with soil depth from surface to deep soil. We concluded that BSCs relieved the extreme hot and cold soil micro-environments in desert ecosystem to some extent. Therefore their effects on soil temperature are positive for improving water and nutrient availability and biological community structure, thus decreasing susceptibility to desertification. These results would be helpful for understanding the ecological and hydrological functions of BSCs in semiarid environment.
Using principal component analysis (PCA), we studied the relationships between hydraulic properties and 13C-CP/MAS-NMR shift regions of biological soil crusts (BSCs) on sand dunes under arid and temperate conditions. The arid study site was located near Nizzana, in the northwestern Negev, Israel and the temperate site was near Lieberose, Brandenburg, Germany. BSCs were sampled at each site near the dune crest, at the center of the dune slope and at the dune base. The soil texture was finer and water holding capacities (WHCs) were higher in Nizzana, whereas surface wettability was reduced in Lieberose. At both sites, BSCs caused extra WHC compared to the mineral substrate. Infiltration after wetting along both catenas generally reached a maximum after 10 min and decreased after 30 min. Carbohydrates were the dominating components in all of the BSCs studied, where the relative peak areas of carbohydrate-derived structures (60–110 ppm) amounted to 28–46% and to 10–14% of total C-peak areas, respectively. PCA revealed that the WHC of the substrate was closely related to the amount of silt and clay, whereas the BSC induced extra WHC was closely related to carbohydrates. It was further found that water repellency was positively related to carbohydrate C, but negatively related to alkyl C. Infiltration kinetics was attributed to polysaccharide hydration and swelling. Our findings support the hypothesis that hydraulic properties of BSCs are determined by extracellular polymeric substances (EPS) and soil texture. Hydraulic properties in BSCs result from the combination of chemical properties related to C compounds mainly dominated by carbohydrates and physical surface properties related to texture, porosity and water holding capacity.
Normalized Difference Vegetation Indices (NDVIs) are typically determined using satellite or airborne remote sensing, or field portable spectrometers, which give an averaged signal on centimetre to metre scale plots. Biological soil crust (BSC) patches may have smaller sizes, and ecophysiological, hydrological as well as pedological processes may be heterogeneously distributed within this level of resolution. A ground-based NDVI imaging procedure using low-cost equipment (Olympus Camedia 5000z digital camera equipped with a Hoya R72 infrared filter) was developed in this study to fill this gap at the level of field research, where carrying costly and bulky equipment to remote locations is often the limiting factor for data collection. Method principle and field data are presented, and the field experiment was deepened comparing NDVI measurements and CO2 turnover of soil crust samples in the laboratory, backing the reliability of the approach.
A commercially available colour rendition chart with known red (600–700 nm) and NIR (800–900 nm) reflectances was placed into each scene and used for calibration purposes on a per-image basis. Generation of NDVI images involved (i) determination of red and NIR reflectances from the pixel values of the red and NIR channels, respectively, and (ii) calculation and imaging of the NDVI, where NDVI values of −1 to +1 were mapped to grey values of 0 to 255. The correlation between NDVI values retrieved from these images and NDVI values determined using field spectrometry was close (r2 = 0.91), the 95% confidence interval amounted to 0.10 NDVI units. The pixel resolution was 0.8 mm in the field and 0.2 mm in the laboratory, but can still be improved significantly with closer distance to the crust or with higher camera resolution. NDVI values obtained using the new method were related to the net CO2 uptake of BSCs, where both slope and correlation coefficient of the respective regression function conformed with literature data. Geostatistical analysis revealed that both spatial variability of net CO2 uptake as well as size of individual hot spots of this parameter increased with crust development. The latter never exceeded 4 mm in the investigated crusts, which points to the necessity of high resolution imaging for linking remote sensing with ecophysiology. Perspectively, the new method could be used for field monitoring of both biological soil crusts and vascular vegetation.
Although biological soil crusts can be found in open landscapes worldwide, their species composition depends on soil properties such as texture and pH, on microclimate, and their respective developmental stage. In addition, local variations in water holding capacity and/or chemical properties of soils influence the formation of spatial patterns and different types of biocrusts on the landscape level. For the evaluation of biocrusts functions and their impact on soil carbon pools, the analysis of the interrelationship between photosynthetic activity and the variations of spatial distribution pattern and types of biocrust is indispensable. For this purpose, an image processing approach was applied that combined chlorophyll fluorescence analyses and multispectral BNDVI to comprehensively characterize the spatial patterns of photosynthetic hotspots in biological soil crusts. For image analysis, five biological soil crust samples with different ratios of substrate, mosses and lichens were collected on an inland dune system in Lieberose, dominated by the moss Polytrichum piliferum, and the lichens Cladonia fimbriata and C. coccifera. RGB-images of the biocrusts were taken with a standard consumer camera Nikon 5200, BNDVI images with a modified Canon S110 NIR camera and chlorophyll fluorescence images with a modular open FluorCAM FC 800-O/1010, respectively. BNDVI and Fv/Fm were nearly in the same range for all biocrust samples related to the total surface area. Although mosses showed higher BNDVI than lichens within the separate biocrust samples. F0 and Fm increased with species coverage and with advancing biocrust development. Overlapping of BNDVI with F0 and Fm images showed that not all crustal organisms contribute to BNDVI and chlorophyll fluorescence. The overlapping areas of BNDVI and F0 ranged between 13% and 29%, that of BNDVI and Fm between 17% and 47%. Matching of RGB, BNDVI and CFI allows visualizing spatial pattern with high or low photosynthesis in biocrusts.
The generally accepted assumption that CAM is an important adaptation to the drought conditions in arid deserts is mainly based on studies in controlled environments comparing the respective photosynthetic pathways in plants of different morphology. Comparative long-term field studies on the ecophysiology of C3- and CAM succulents co-occurring in the same desert habitat are too rare to satisfactorily answer the old question whether the metabolic pathway or the succulence is more important to guarantee survival in such a harsh habitat. Thus, relevant ecophysiological parameters were repeatedly monitored on plants of the leaf succulents Augea capensis (C3) and Malephora purpureo-crocea (CAM), which commonly dominate a habitat in the Knersvlakte, Namaqualand, South Africa. Under identical conditions in the same habitat, carbon gain of the C3 species was higher than that of the CAM plant, but their WUE lower and maximum photosynthetic activity nearly the same. In M. purpureo-crocea, irrigation induced rapid whole day stomatal opening at drastically increased daytime but decreased night-time CO2-uptake, while respective responses of the C3 species were delayed. These different response dynamics were due to different rooting depth, root morphology and water acquisition strategy of the flat rooting CAM and the deep-rooting C3 species. This, however, may not explain the highly similar growth success. Anyway, the results verify that CAM provides no advantage over C3 for long-term survival in the same desert environment in morphologically similar succulents.
In nutrient-poor ecosystems, certain plant families have evolved specific adaptations for phosphate acquisition, namely cluster roots or proteoid roots. Cluster rooted species have a unique suite of adaptations that enable them to survive in nutrient poor ecosystems, such as the Fynbos biome of South Africa. The Fynbos biome has heterogeneous soils which caused radiation of varying soil pH within small localised areas. Previous studies on cluster rooted species ignored the possibility that cluster root functioning may differ at varying soil pH levels and soil types. Therefore, two native Fynbos protea species, Protea cynaroides (L.) L. and Protea obtusifolia H.Buek ex Meisn. were compared to assess the impact of different soil pH levels on organic acid metabolism and exudation in cluster rooted species. Our results show that cluster roots of the acidic-soil grown P. cynaroides is more effective and more efficient than that of the alkaline-soil grown P. obtusifolia. Our findings indicate that the mechanisms for P acquisition vary greatly, depending on substrate and plant organ type. This suggests that the metabolism of each species is adapted to specific soil pH and that these species have evolutionary adapted their P acquisition mechanisms for these extreme soils. The limited nutrient availability and variation in soil pH South African soils has led to various localised adaptations for cluster rooted species.
Under the conditions of climate change in South Africa, ecological and technical measures are needed to reduce the water consumption of irrigated crops. Windbreak hedges are long-rated systems in agriculture that significantly reduce wind speed. Their possibilities to reduce evapotranspiration and water demand are being investigated at a vineyard in the Western Cape Province, South Africa. Detailed measurements of meteorological parameters relevant for the computation of reference and crop-specific evapotranspiration following the FAO 56 approaches within a vineyard in the Western Cape Province of South Africa have shown the beneficial effect of an existing hedgerow consisting of 6 m high poplars (Populus simonii (Carrière) Wesm.). With reference to a control station in the open field, the mean wind speed in a position about 18 m from the hedgerow at canopy level (2 m) was reduced by 27.6% over the entire year and by 39.2% over the summer growing season. This effect leads to a parallel reduction of reference evapotranspiration of 15.5% during the whole year and of 18.4% over the growing season. When applying empirical crop-specific Kc values for well-irrigated grapes, the reduction of evapotranspiration is 18.8% over the summer growth period. The introduced tree shelterbelts are a suitable eco-engineering approach to reduce water consumption and to enhance water saving in vineyards.
It is commonly accepted that CO2 efflux increases with soil water content in aerated soils and that rewetting after periods of soil drying can result in respiration pulses. It has further been shown that soil pores may become water logged which can impede soil gas exchange. The present study aimed to quantify the carbon response of moss (Polytrichum piliferum) and lichen (Cladonia coniocraea) biocoenoses to different quantities of rain in an artificial catchment, which granted that the starting point of the development of both biocoenoses was the same. To address this aim, we conducted in situ soil moisture and soil respiration measurements, where soil respiration was hypothesised to emerge from a cryptogamic vegetation layer and from mineral soil beneath. We found that higher water-holding capacity of the moss layer and higher accumulation of organic matter in the upper mineral soil under mosses result in higher amounts of water stored near the surface. As a consequence, evaporation of water as well as pulses of CO2 efflux after moderate rain following a period of drought were higher in the moss biocoenosis, where the upper mineral soil was of key importance. In contrast, the lichen biocoenosis facilitated penetration of rainwater into the deep soil. Superimposing rewetting pulses, near-saturation of soil pores with water after severe rain resulted in gas exchange inhibition and diminished soil respiration until subsequent aeration in both biocoenoses
Windbreaks, trees, structures, and shade nets are used in South Africa to protect citrus orchards from wind scar damage to avoid major losses in export quality. Severe southeasterly winds in the Western Cape during spring and summer are the main causes of significant external peel damage. In this study, wind scar damage of ‘Tango’ mandarin (Citrus reticulata) fruit was quantified at different distances from established Casuarina cunninghamiana (Beefwood) and Populus simonii (Chinese Poplar) windbreaks around Stellenbosch, Western Cape, South Africa. The yield efficiency was not significantly affected by distance from the windbreak in either of the trials. Fruit at 2 H and 3 H had significantly lower ratios of total soluble solids to titratable acidity, but export standards for mandarins were still met. The wind speed was consistently higher at 16 H, compared with 3 H during summer. The highest packout (class 1 fruit) was observed near the windbreak (3 H) and decreased as the distance increased towards 10 H and 16 H. There were no significant differences for severe wind scar damage incidence between the distance treatments. Our results confirmed the impact of wind on external fruit quality and the limitation of tree-based windbreaks with an increase in distance, which partly supports the current movement towards full enclosure of orchards with shade nets.
Als mehrjährige Bioenergiepflanze kann Sida hermaphrodita eine wichtige Rolle für den Anbau auf marginalen Standorten in der nahen Zukunft spielen. In einem kontrollierten Topfexperiment wurde der Einfluss von verschiedenen Kompostgehalten (10, 20, 30, 50%) und mineralischem Stickstoffdünger (100 kg N ha–1) auf Biomasse, Wuchshöhe und Kohlenstoffallokation während der Etablierungsphase untersucht. Weiterhin wurden ökophysiologische Parameter (Gaswechsel, Chlorophyllfluoreszenz, Chlorophyllgehalte) mit nicht-destruktiven Methoden bestimmt. Die Ergebnisse des Experimentes zeigen, dass sowohl die Stickstoffdüngung als auch die Kompostgabe positive Effekte für das Wachstumsverhalten hat. Während der Etablierungsphase erreichten die gedüngten Pflanzen eine mittlere Pflanzenhöhe von bis zu 127 cm (maximale Wuchshöhe 168 cm). Die Photosynthese wurde nicht durch die unterschiedlichen Behandlungen beeinflusst. Die mittlere CO2-Austauschrate variiert zwischen 5,8 und 9,9 μmol m–2 s–1 und die Elektronentransportrate zwischen 34,5 und 53,9 μmol m–2 s–1. Die Verwendung von Kompost und Stickstoff ist für eine bessere Biomasseproduktion von Sida in marginalen Böden möglich, auch wenn das Wachstum im ersten Jahr limitiert ist und es negative Rückwirkungen auf die Bestandesetablierung gibt.
Trees from drought deciduous forests (“bosque seco”) in northern Peru and southern Ecuador regularly shed their leaves at the onset of seven months-long periods without rain. This way they very effectively reduce the bulk water consumption by leaf transpiration, still loosing small amounts of water through their bark, but at the risk of too little or no carbon gain when leaf-less. Seven species studied here have good developed green cortex tissue on stems and axes, as especially evident for the “bottle tree” Ceiba trischistandra (A. Gray) Bakhuisen (Bombacaceae). In several leaf-less tree species at the end of summer drought, when xylem sap flows are very low, indications of higher xylem flows at night as compared to day-time were found, which led to the question of whether the trees are capable of performing CAM in their axes. However, although quite low in some cases, the pH of the cortex tissues did not fluctuate diurnally. In addition, no night-time carbon uptake could be observed. Although no net carbon gain was measured in C. trischistandra and Erythrina smithiana Krukoff (Fabaceae) photosynthetic activity of the green cortex was sufficient to re-assimilate between 50% to 60% of the carbon released by mitochondrial respiration. In contrast, the obligate CAM plant Cereus diffusus (Britton & Rose) Werdermann (Cactaceae), which was studied as a reference in the same environment, showed both diurnal pH-fluctuations in its green tissue with lowest values before sun rise, and net carbon fixation at night.
Cyanobacteria and green algae present in biological soil crusts are able to colonize mineral substrates even under extreme environmental conditions. As pioneer organisms, they play a key role during the first phases of habitat colonization. A characteristic crust was sampled 3 years after installation of the artificial water catchment “Chicken creek”, thus representing an early successional stage of ecosystem development. Mean annual rainfall and temperature were 559 mm and 9.3°C, respectively. We combined scanning electron microscopy (SEM/EDX) and infrared (FTIR) microscopy to study the contact zone of algal and cyanobacterial mucilage with soil minerals in an undisturbed biological soil crust and in the subjacent sandy substrate. The crust was characterized by an approximately 50 μm thick surface layer, where microorganisms resided and where mineral deposition was trapped, and by an approximately 2.5 mm thick lower crust where mineral particles were stabilized by organo-mineral structures. SEM/EDX microscopy was used to determine the spatial distribution of elements, organic compounds and minerals were identified using FTIR microscopy and X-ray diffraction (XRD). The concentration of organic carbon in the crust was about twice as much as in the parent material. Depletion of Fe, Al and Mn in the lower crust and in the subjacent 5 mm compared to the geological substrate was observed. This could be interpreted as the initial phase of podzolization. Existence of bridging structures between mineral particles of the lower crust, containing phyllosilicates, Fe compounds and organic matter (OM), may indicate the formation of organo-mineral associations. pH decreased from 8.1 in the original substrate to 5.1 on the crust surface 3 years after construction, pointing to rapid weathering of carbonates. Weathering of silicates could not be detected.
Dew formation was investigated in three developmental stages of biological soil crusts (BSC), which were collected along a catena of an inland dune and in the initial substrate. The Penman equation, which was developed for saturated surfaces, was modified for unsaturated surfaces and used for prediction of dewfall rates. The levels of surface saturation required for this approach were predicted using the water retention functions and the thicknesses of the BSCs. During a single event, dewfall increased with crust development from 0.08 kg m−2 for the initial substrate to 0.10, 0.20 and 0.25 kg m−2 for crusts stages 1 to 3, respectively, which was well reflected by the model response. The suggested mechanism of dew formation involves a delay in water saturation in near-surface soil pores and EPS where the crusts were thicker and where the water capacity was high, resulting in elevated vapor flux towards the surface. The results also indicate that the amount of dewfall was too low to observe water flow into deeper soil. Analysis of the soil water retention curves revealed that, despite the sandy mineral matrix, moist crusts with clogged by swollen EPS pores exhibited a clay-like behavior. It is hypothesized that BSCs gain double benefit from suppressing their competitors by runoff generation and from improving their water supply by dew collection. Despite higher amounts of dew, the water availability to the crust community decreases with crust development, which may be compensated by ecophysiological adaptation of crust organisms, and which may further suppress higher vegetation or mosses.
The influence of biological soil crusts in natural ecosystems on structures and processes is well investigated. However, in South-Brandenburg (Germany), it is possible to study the development of biological soil crusts (BSC) during initial ecosystem genesis on two artificial water catchments with well-known ages and under differing starting conditions. The two experimental sites are located in the recultivation area of the lignite open-cast mining district of southern Brandenburg with a distance of approximately 1 km between them. Two different topographies were constructed at the experimental sites: the experimental plot at the catchment Neuer Lugteich was shaped like a dune, whereas the artificial water catchment Hühnerwasser was modelled as an inclined slope. The catchment Neuer Lugteich is four years older than Hühnerwasser. The original substrate at Neuer Lugteich is more sandy and carbonate-free compared to the original substrate dumped at Hühnerwasser. At both sites geomorphological differentiation and crust development were compared and the importance of substrate-dependent water availability and crust type clarified. Once settled, the crusts influenced the water regime of the soils by delaying infiltration through enhanced water repellency, and by limiting water infiltration. Chlorophyll analysis revealed that all crusts were at early stages of development. At Neuer Lugteich, the establishment of the biological soil crusts was closely associated with the vegetation succession, whereas no clear succession of the crusts could be observed at Hühnerwasser. The mosaic-like pattern of the biological soils crusts is associated with the distribution of fine-grained material here.zeige weniger