Bingchang Zhang

Microalgal species variation at different successional stages in biological soil crusts of the Gurbantunggut Desert, Northwestern China

Biological soil crusts (BSC), most notably lichen crusts, develop and diversify in the Gurbantunggut Desert, the largest fixed and semi-fixed desert in China. Four different successional stages of BSC, including bare sand, microalgal crusts, lichen crusts, and moss crusts, were selected to determine successional changes in microalgal species composition and biomass and formation of BSC. A 10 × 10-m observation plot was established in an interdune region of the Gurbantunggut Desert and data were collected over an 8-year study period. The main results were: (1) different successional stages of BSC significantly affected the content of soil organic C and total and available N but not the total and available P and K content of soil; (2) composition of microalgal communities differed among the four successional stages; (3) significant differences in microalgal biomass were observed among the four successional stages; (4) bare sand was mainly uncompacted sand gains; (5) filamentous cyanobacteria, particularly Microcoleus vaginatus, were the dominant species in the early phase of crust succession. The presence of fungal mycelium and moss rhizoids prevented water and wind erosion.

Species composition,distribution patterns and ecological functions of biological soil crusts in the Gurbantunggut Desert

As one of the most important biological factors that maintain the stability of the largest fixed and semi-fixed desert in China,the Gurbantunggut Desert,the biological soil crusts (BSCs) develop well and play critical ecological roles in the desert ecosystem. In this paper,we briefly summarize our research findings since 2002 including species composition,distribution pattern and ecological functions of BSCs in the desert. Our results indicate abundant species diversity of BSCs in the Gurbantunggut Desert in comparison to other deserts in China. At the scales of sand dune or whole desert,the distribution patterns of BSCs are location-specific. The existence of BSCs in this desert could:(1) accelerate the formation of desert soil and the weathering of minerals; (2) accumulate organic matter in surface soil through related species in soil crusts; (3) enhance the abilities of sand surface to resist wind erosion; (4) influence seed germination of vascular plants; and (5) enhance the production of dew deposition on sandy soil surface.

Distribution and Composition of Cyanobacteria and Microalgae Associated with Biological Soil Crusts in the Gurbantunggut Desert, China

In Gurbantunggut Desert, cyanobacterial and microalgal components were characterized within 60 soil samples collected from sand dunes. Fifty-one taxa of cyanobacteria and algae were identified. Without exception, the soils were alkaline, poor in nutrients, and showed large variations in other soil properties. Spatial heterogeneity for distribution of cyanobacteria and microalgae (diversity of morphotypes, species composition, and microbiomass) existed. Compared with other deserts in the world, the Gurbantunggut Desert has a greater diversity of cyanobacterial-microalgal morphotypes. Results from step regression showed that the diversity of morphotype was determined by total P, available P, and soil layer. Filamentous cyanobacteria dominated the community. Microcoleus vaginatus (Vauch.) Gom was the dominant species in most positions on sand dune, while the abundance of other dominant species varied depending on the sand dune position and the soil layer in which they occurred. The microalgal biomass was influenced by the content of Mg, crust type, soil moisture, sunlight, and oxygen concentration. A significant positive relation was found between microalgal biomass and diversity of morphotype. Species composition, diversity of morphotype, and microalgal biomass interacted with each other. The contents of P and Mg ion, soil texture, and soil moisture may be the main factors responsible for cyanobacterial-microalgal distribution.

Photosynthetic and cytological recovery on remoistening Syntrichia caninervis Mitt., a desiccation-tolerant moss from Northwestern China

Syntrichia caninervis Mitt. is the dominant species in the moss crusts of the Gurbantunggut Desert, Northwestern China. We experimented with this species under controlled environmental conditions. Modulated chlorophyll (Chl) fluorescence was used to test the speed of recovery as evidenced by the time course of photosynthetic activity following remoistening. Transmission electron microscopy was used to explore the cytological characteristics of the leaf cells. Minimum and maximum fluorescence (F0 and FM) and photosynthetic yield (FV/FM) of photosystem II (PSII) recovered quickly when shoots were remoistened in the dark. This was especially the case of FV/FM; within the first minute of remoistening this reached 90% or more of the value attained after 30 min. These physiological changes were closely paralleled by cytological changes that indicated no damage to membranes or organelles. Correlation analysis showed that Chl fluorescence decreased both above and below a narrow moisture optimum. Our results underline the capability of S. caninervis to photosynthesize after remoistening. Utilizing precipitation events such as dew, fog, rain, and melting snow allows S. caninervis to survive and grow in a harsh desert environment.

Morphological Adaptations to Drought and Reproductive Strategy of the Moss Syntrichia caninervis in the Gurbantunggut Desert, China

The biological soil crusts (BSCs) in the Gurbantunggut Desert, the largest fixed and semi-fixed desert in China, feature moss-dominated BSCs, which play an indispensable role in sand fixation. Syntrichia caninervis Mitt. (S. caninervis) serves as one of the most common species in BSCs in the desert. In this study we examined the morphological structure of S. caninervis from leafy gametophyte to protonema using light and scanning electron microscopy (SEM). We also examined the relationships between the morphological structure of S. caninervis and environmental factors. We found that: (1) this moss species is commonly tufted on the sand surface, and its leaves are folded upwards and twisted around the stem under dry conditions; (2) the cells on both upper and lower leaf surfaces have C-shaped dark papillae, which may reflect sunlight to reduce the damage from high temperature; (3) the leaf costa is excurrent, forming an awn with forked teeth; and (4) the protonema cells are small and thickset with thick cell walls and the cytoplasm is highly concentrated with a small vacuole. In addition, we also found that the protonema cells always form pouches on the tip of the mother cells during the process of cell polarization. Our results suggest that S. caninervis has, through its life cycle, several morphological and structural characteristics to adapt to dry environmental conditions. These morphological features of S. caninervis may also be found in other deserts in the world due to the world-wide distribution of the species.

Bacterial diversity and community along the succession of biological soil crusts in the Gurbantunggut Desert, Northern China.

Biological soil crusts (BSCs) are common and play critical roles in semi‐arid and arid ecosystems. Bacteria, as an important community in BSCs, play critical roles in biochemical processes. However, how bacterial diversity and community change in different successional stages of BSCs is still unknown. We used 454 pyrosequencing of 16S rRNA to investigate the bacterial composition and community, and the relationships between bacterial composition and environmental factors were also explored. In different successional stages of BSCs, the number of bacteria operational taxonomic units (OTUs) detected in each sample ranged from 2572 to 3157. Proteobacteria, Cyanobacteria, Bacteroidetes were dominant in BSCs, followed by Firmicutes, Acidobacteria, and Actinobacteria. At the successional stages of BSCs, bacterial communities, OTU composition and their relative abundance notably differentiated, and Cyanobacteria, especially Microcoleus vaginatus, dominated algal crust and lichen crust, and were the main C‐fixing bacteria in BSCs. Proteobacteria and Bacteroidetes increased with the development of BSCs. OTUs related to Planomicrobium Chinese, Desulfobulbus sp., Desulfomicrobium sp., Arthrobacter sp., and Ahhaerbacter sp. showed higher relative abundance in bare sand than other successional stages of BSCs, while relative abundance of Sphingomonas sp. Niastella sp., Pedobacter, Candidatus solobacter, and Streptophyta increased with the development of BSCs. In successional stages of BSCs, bacterial OTUs composition demonstrated strong correlations with soil nutrients, soil salts, and soil enzymes. Additionally, variation of bacterial composition led to different ecological function. In bare sand, some species were related with mineral metabolism or promoting plant growth, and in algal crust and lichen crust, C‐fixing bacteria increased and accumulated C to the desert soil. In later developed stage of BSCs, bacteria related with decomposition of organic matter, such as Sphingomonas sp. Niastella sp., Pedobacter, and Candidatus solobacter increased. Therefore, bacterial community composition and their key ecological roles shifted to the development of BSCs.

Responses of microbial activities and soil physical-chemical properties to the successional process of biological soil crusts in the Gurbantunggut Desert, Xinjiang

Biological soil crusts (BSCs) are capable of modifying nutrient availability to favor the establishment of biogeochemical cycles. Microbial activities serve as critical roles for both carbon and nutrient transformation in BSCs. However, little is known about microbial activities and physical-chemical properties of BSCs in the Gurbantunggut Desert, Xinjiang, China. In the present research, a sampling line with 1-m wide and 20-m long was set up in each of five typical interdune areas selected randomly in the Gurbantunggut Desert. Within each sampling line, samples of bare sand sheet, algal crusts, lichen crusts and moss crusts were randomly collected at the depth of 0–2 cm. Variations of microalgal biomass, microbial biomass, enzyme activities and soil physical-chemical properties in different succession of BSCs were analyzed. The relationships between microalgal biomass, microbial biomass, enzymatic activities and soil physical-chemical properties were explored by stepwise regression. Our results indicate that microalgal biomass, microbial biomass and most of enzyme activities increased as the BSCs developed and their highest values occurred in lichen or moss crusts. Except for total K, the contents of most soil nutrients (organic C, total N, total P, available N, available P and available K) were the lowest in the bare sand sheet and significantly increased with the BSCs development, reaching their highest values in moss crusts. However, pH values significantly decreased as the BSCs developed. Significant and positive correlations were observed between chlorophyll a and microbial biomass C. Total P and N were positively associated with chlorophyll a and microbial biomass C, whereas there was a significant and negative correlation between microbial biomass and available P. The growth of cyanobacteria and microorganism contributed C and N in the soil, which offered substrates for enzyme activities thus increasing enzyme activities. Probably, improvement in enzyme activities increased soil fertility and promoted the growth of cyanobacteria, eukaryotic algae and heterotrophic microorganism, with the accelerating succession of BSCs. The present research found that microalgal-microbial biomass and enzyme activities played important roles on the contents of nutrients in the successional stages of BSCs and helped us to understand developmental mechanism in the succession of BSCs.

Responses of microalgal-microbial biomass and enzyme activities of biological soil crusts to moisture and inoculated Microcoleus vaginatus gradients.

Microcoleus vaginatus Gom., the dominant cyanobacterial species in biological soil crusts (BSCs), plays a critical role in forming BSCs. Four inoculum dosages of M. vaginatus (0.5, 1, 3, and 7 g · m−2) were inoculated across a sand surface and subjected to five moisture treatments (0, 1, 2, 3, and 4 L · m−2). Microalgal biomass, microbial biomass, and enzyme activities (invertase, alkaline phosphatase, and urease) were subsequently determined. In general, microalgal biomass, microbial biomass composed of carbon and nitrogen, as well as alkaline phosphatase and urease activity were increased by increasing inoculum dosages and moisture, whereas invertase activity decreased with higher moisture content. Combination of 7 g m−2 inoculum and 2 or 3 L m−2 moisture was recommended for M. vaginatus to recover BSCs. Significant positive correlations were observed between microalgal biomass, microbial biomass, C:N ratios, and enzyme activities. Increasing microalgal biomass can act as an energy source for microbial activity, while the growth of cyanobacteria and microorganisms provides substrates for enzymatic activities. Results from the present study allows better understanding of the conditions for M. vaginatus forming BSCs and offers guidance for successful recovery of BSCs in the field.

Cyanobacterial composition and spatial distribution based on pyrosequencing data in the Gurbantunggut Desert, Northwestern China

Cyanobacteria are the primary colonizers and form a dominant component of soil photosynthetic communities in biological soil crusts. They are crucial in improving soil environments, namely accumulating soil carbon and nitrogen. Many classical studies have examined cyanobacterial diversity in desert crusts, but relatively few comprehensive molecular surveys have been conducted. We used 454 pyrosequencing of 16S rRNA to investigate cyanobacterial composition and distribution on regional scales in the Gurbantunggut Desert. The relationship between cyanobacterial distribution and environmental factors was also explored. A total of 24,973 cyanobacteria partial 16S rRNA gene sequences were obtained, and 507OTUs were selected, as most OTUs had very few reads. Among these, 347 OTU sequences were of cyanobacteria origin, belonging to Oscillatoriales, Nostocales, Chroococcales, and uncultured cyanobacterium clone, respectively. Microcoleus vaginatus, Chroococcidiopsis spp. and M. steenstrupii were the dominant species in most areas of the Gurbantunggut Desert. Compared with other desert, the Gurbantunggut Desert differed in the prominence of Chroococcidiopsis spp. and lack of Pseudanabaenales. Species composition and abundance of cyanobacteria also showed distinct variations. Soil texture, precipitation, and nutrients and salt levels affected cyanobacterial distribution. Increased precipitation was helpful in improving cyanobacterial diversity. A higher content of coarse sand promoted the colonization and growth of Oscillatoriales and some phylotypes of Chroococcales. The fine‐textured soil with higher nutrients and salts supported more varied populations of cyanobacteria, namely some heterocystous cyanobacteria. The results suggested that the Gurbantunggut Desert was rich in cyanobacteria and that precipitation was a primary regulating factor for cyanobacterial composition on a regional scale.

Progress in the study of algae and mosses in biological soil crusts

Algae and mosses are not only two of the familiar communities in the process of desert vegetational succession, but also have the highest biomass in biological soil crusts. Meanwhile, being the pioneer plants, algae and mosses are involved in the establishment of biological soil crusts, which have great importance in arid environments and play a major role in desert ecosystems, such as being the indicator of the vegetation type, soil-holding, preventing erosion by water and wind, and sand fixation. This paper reviews the advances in the study of algae and mosses in arid and semi-arid areas. It mainly describes the ecological functions of algae and mosses including their influences on water cycle, circulation of substances, and community succession. In addition, the relationships between algae and mosses are discussed. Finally, some suggestions are proposed for the research orientations of algae and mosses in biological soil crusts. Ecologically, algae and mosses have significant ecological importance in arid areas, especially in those areas where environmental problems are becoming increasingly serious.