Interspecific root interaction enhances cadmium accumulation in Oryza sativa when intercropping with cadmium accumulator Artemisia argyi.

The contamination of arable land with heavy metals, such as Cd, is a serious concern worldwide. Intercropping with Cd accumulators can be used for efficient safe crop production and phytoremediation of Cd-contaminated soil. However, the effect of intercropping on Cd uptake by main crops and accumulators varies among plant combinations. Rhizosphere interaction may mediate Cd uptake by intercropped plants, but the mechanism is unclear. Thus, in the present study, we aimed to examine the effect of rhizosphere interaction on Cd uptake by intercropping rice (Oryza sativa L.) with mugwort (Artemisia argyi Levl. et Vant.) in Cd-contaminated paddy soil. We grew O. sativa and A. argyi in pots designed to allow different levels of interaction: complete root interaction (no barrier), partial root interaction (nylon mesh barrier), and no root interaction (plastic film barrier). Our results indicated that both complete and partial root interaction increased the shoot and root mass of A. argyi, but did not decrease the shoot, root, and grain mass of O. sativa. Interspecific root interaction significantly increased the Cd content in the shoots, roots, and grains of O. sativa and the shoots of A. argyi. Increased content of total organic acids in the rhizosphere, which increased the content of available Cd, was a possible mechanism of increased Cd uptake in both plants under interspecific root interaction. Our findings demonstrate that an intercropping system can extract more Cd from contaminated soil than a monocropping system of either A. argyi or O. sativa. However, the intercropping system did not facilitate safe crop production because it substantially increased grain Cd content in O. sativa.

Potential risk of eutrophication in the deepest lake of Southwest China: Insights from phosphorus enrichment in bottom water.

Large deep lakes in plateau regions provide crucial ecosystem services but are susceptible to eutrophication due to their long water residence time. To date, the water quality of deep lakes has not received as much attention as that of shallow lakes owing to logistical challenges. This study investigated the seasonal variation and vertical distribution of phosphorus and related environmental variables in a large deep lake in the Yunnan Plateau, China (Fuxian Lake). Generally, the concentrations of total phosphorus (TP, R2 = 0.862), total dissolved phosphorus (TDP, R2 = 0.922), and dissolved inorganic phosphorus (DIP, R2 = 0.889) exhibited a linear increase with the greater water depth, whereas the pH and dissolved oxygen (DO) showed decreasing trends. The TP, TDP, and DIP values were 0.012, 0.006, and 0.004 mg/L, respectively, in surface waters (0.5 m depth), and increased to 0.074, 0.065, and 0.062 mg/L, respectively, at 140.0 m depth. The averaged over ordering method demonstrated that DO and air temperature accounted for a higher proportion of the explained variance of TP, TDP, and DIP in the shallow water layer (0.5-20.0 m). In contrast, DO and pH accounted for a higher proportion of the explained variance of TP, TDP, and DIP in deeper water layers (40.0-150.0 m). As a warm monomictic lake, the higher observed phosphorus concentrations in deeper water and sediment potentially pose a risk of future eutrophication in the Fuxian Lake. Our findings demonstrate that more efficient technical and management measures should be taken to reduce the external phosphorus load to Fuxian Lake, so that the load to and from the sediment will decrease eventually.

Mapping Chinese annual gross primary productivity with eddy covariance measurements and machine learning.

Annual gross primary productivity (AGPP) is the basis for grain production and terrestrial carbon sequestration. Mapping regional AGPP from site measurements provides methodological support for analysing AGPP spatiotemporal variations thereby ensures regional food security and mitigates climate change. Based on 641 site-year eddy covariance measuring AGPP from China, we built an AGPP mapping scheme based on its formation and selected the optimal mapping way, which was conducted through analysing the predicting performances of divergent mapping tools, variable combinations, and mapping approaches in predicting observed AGPP variations. The reasonability of the selected optimal scheme was confirmed by assessing the consistency between its generating AGPP and previous products in spatiotemporal variations and total amount. Random forest regression tree explained 85 % of observed AGPP variations, outperforming other machine learning algorithms and classical statistical methods. Variable combinations containing climate, soil, and biological factors showed superior performance to other variable combinations. Mapping AGPP through predicting AGPP per leaf area (PAGPP) explained 86 % of AGPP variations, which was superior to other approaches. The optimal scheme was thus using a random forest regression tree, combining climate, soil, and biological variables, and predicting PAGPP. The optimal scheme generating AGPP of Chinese terrestrial ecosystems decreased from southeast to northwest, which was highly consistent with previous products. The interannual trend and interannual variation of our generating AGPP showed a decreasing trend from east to west and from southeast to northwest, respectively, which was consistent with data-oriented products. The mean total amount of generated AGPP was 7.03 ± 0.45 PgC yr-1 falling into the range of previous works. Considering the consistency between the generated AGPP and previous products, our optimal mapping way was suitable for mapping AGPP from site measurements. Our results provided a methodological support for mapping regional AGPP and other fluxes.

Concurrent effects of flooding regimes and floodwater quality on sediment properties in a Yangtze River-connected floodplain wetland: Insights from field investigations during 2011-2020.

Changes in flood regimes, floodwater quality, and macrophyte types may affect sediment characteristics post-flooding. However, few studies have attempted to unravel their complex influences in floodplain wetlands. From 2011 to 2020, the physical and chemical properties of surface layer sediment pre- and post-flooding was investigated through field surveys in the Dongting Lake wetland. Results indicated that the pre-flooding soil total phosphorus (TP) and total nitrogen (TN) exhibited an increasing trend during 2011-2020. Soil TP increased post-flooding relative to that pre-flooding. The changes in TN, sediment organic matter (SOM), sediment moisture content (SMC), and sediment bulk density (SBD) fluctuated over the years. The best-fitting multi-regression model demonstrated that the changes in sediment variables post-flooding showed a parabolic trajectory along the inundation duration (ID), except for SMC. Changes in soil properties post-flooding were negatively correlated with ID for sediment with a low IDs (<148 days). Meanwhile, changes in soil properties post flooding were positively correlated with ID for sediment with a high IDs (>193 days). Changes in SBD and SOM post-flooding were positively influenced by the TP content in the floodwater. These findings indicate that changes in the flooding regime, and water quality generated by anthropogenic disturbances such as the Three Gorges Dam significantly affect sediment properties, and subsequently influence the ecological functions of the Dongting Lake wetland.

Belowground seed and bud banks play complementary roles in the potential recruitment of dominant macrophyte communities in a Yangtze River-connected floodplain wetland.

Both seed and bud banks play important roles in the recruitment and maintenance of macrophyte communities; however, few studies have investigated them simultaneously. We investigated the abundance, species composition, and seasonal patterns of seed and bud banks in two dominant macrophyte communities, Carex and Miscanthus, in the Dongting Lake wetlands. The seed densities of both communities were lower from November (after flooding) to March and increased dramatically before flooding (in May). The bud densities of the two dominant communities peaked in the coldest month of the year (January), decreased markedly in March, and were the lowest before flooding. The seed banks of the two macrophyte communities were mainly composed of annual species and a few perennial species, whereas the bud banks were composed of only dominant perennials. Furthermore, the perennial species present in bud banks did not occur in seed banks. Among the soil variables, the bud densities of both plant communities were negatively associated with soil bulk density, whereas the seed density of the Miscanthus community was positively associated with soil bulk density. Our results suggest that seed and bud banks are complementary in the potential recruitment of macrophyte communities; that is, bud banks regulate the demography of dominant perennials, and seed banks contribute to the recruitment and dispersal of annual species. Given the high abundance of annuals and near absence of the most dominant perennials in the seed bank, the bud banks of dominant perennial species should be more widely used in wetland restoration and management.

Exploring the relative contribution of flood regimes and climatic factors to Carex phenology in a Yangtze River-connected floodplain wetland.

Hydrological regimes can combine with climatic factors to affect plant phenology; however, few studies have attempted to quantify their complex influences on plant phenology in floodplain wetlands. We obtained phenological information on Carex vegetation through MODIS normalized difference vegetation index (NDVI) data during 2001-2020, and monthly field investigation during 2011-2020. We then explored how these data were correlated with climatic factors and flood regimes in a Yangtze River-connected floodplain wetland (Dongting Lake, China). Our results showed that warmer temperature tended to advance the start of the pre-flooding growing season (SOS1), with a relative contribution of 76.1 %. Flood rising time strongly contributed to controlling the end of the pre-flooding growing season. Flood recession time and inundation duration were dominant factors determining the start of the post-flooding growing season (SOS2). Earlier flood recession time and shortened inundation duration tended to advance the SOS2. Shortened inundation duration, earlier flood recession time, and lower solar radiation tended to advance the end of the post-flooding growing season. The phenology of Carex distributed at high-elevation areas was more affected by hydrology than that of Carex distributed at low-elevation areas. Thus, climatic factors strongly affect the phenology of Carex during the pre-flooding growing season, whereas flood regimes play a dominant role in determining the phenology in the post-flooding growing season. The different responses of Carex phenology to climatic and flooding factors may provide insights for the conservation and management of floodplain wetlands in Yangtze River because Carex are primary food source and habitat for herbivorous waterfowls.

Community Trait Responses of Three Dominant Macrophytes to Variations in Flooding During 2011-2019 in a Yangtze River-Connected Floodplain Wetland (Dongting Lake, China).

In lacustrine wetlands connected to rivers, the changes in flood regimes caused by hydrological projects lead to changes in the community traits of dominant macrophytes and, consequently, influence the structure and function of wetland vegetation. However, community trait responses of macrophytes to the timing and duration of flood disturbance have been rarely quantified. In 2011-2019, we investigated plant species diversity, density, and biomass in three dominant macrophyte communities (Carex brevicuspis C.B. Clarke, Miscanthus sacchariflorus (Maxim.) Hackel, and Polygonum hydropiper L.) through monthly field surveys in Dongting Lake wetlands. Partial least squares regressions were used to analyze how the variations in hydrological regimes affected plant community traits. Apparent inter-annual fluctuations in plant community traits were detected during 2011-2019. The species richness and Shannon index of diversity of Miscanthus and Polygonum communities increased, whereas the Shannon index of diversity of Carex community decreased. Variation in flooding had a greater effect on Polygonum and Carex community traits than on Miscanthus community traits. Flooding disturbed all plant communities, especially when the duration and timing varied. Shorter inundation periods caused the biomass of Miscanthus community to decline, and that of Carex and Polygonum communities to increase. Earlier flood recession caused the species richness and Shannon index of diversity of Polygonum and Miscanthus community to increase, and those of Carex community to decrease. These findings imply that shorter inundation durations and earlier flood recession generated by the operation of the Three Gorges Dam have changed the macrophyte growth pattern.

Testing the Growth Rate Hypothesis in Two Wetland Macrophytes Under Different Water Level and Sediment Type Conditions.

The growth rate hypothesis (GRH) states that a negative correlation exists between the growth rate and N:P and C:P ratios, because fast-growing organisms need relatively more phosphorus-rich RNA to support their high rates of protein synthesis. However, it is still uncertain whether the GRH is applicable in freshwater wetlands. Several studies have shown that water level and sediment type are key factors influencing plant growth and plant C:N:P characteristics in freshwater wetlands. Thus, this study aimed to elucidate the influence of these factors on plant growth and test the GRH under varying water levels and sediment conditions. We designed a controlled experiment at three water levels and under three sediment types using the two dominant plants (Carex brevicuspis and Polygonum hydropiper) in the East Dongting Lake wetland, and we further investigated the relative growth rate (RGR); concentrations of total carbon (TC), total nitrogen (TN), and total phosphorus (TP); and plant stoichiometry (ratios of C:N, C:P, and N:P) in the aboveground and belowground parts and whole plants in both species. Results demonstrated that the RGR and TC of both species decreased significantly with decreasing sediment nutrient supply and increasing water level. However, TN and TP of both species were markedly higher at high water levels than at low water levels; furthermore, these were significantly higher on clay than on the other two sediment types at each water level. The C:N and C:P ratios of both species decreased with increasing sediment nutrient supply and water level, whereas N:P decreased in both species with increasing sediment nutrient supply. The aboveground part of C. brevicuspis as well as the aboveground part and whole plant of P. hydropiper were negatively correlated with N:P, which is consistent with the GRH. However, the relationship between the belowground RGR and N:P of these species was inconsistent with GRH. Therefore, the water level and sediment type and their interaction significantly influenced plant RGR and C:N:P characteristics. The RGR and plant stoichiometry differed significantly between plant organs, indicating that the GRH needs refinement when applied to wetland macrophytes.

Crucial sites and environmental variables for wintering migratory waterbird population distributions in the natural wetlands in East Dongting Lake, China.

Dongting Lake is the second largest freshwater lake in China and is one of the globally important wintering sites for migratory waterbirds in the East Asian-Australasian Flyway. Crucial sites and environmental variables for wintering migratory waterbirds are of great concern in the Dongting Lake wetlands. In this research, based on annual (2003/2004-2016/2017) waterbird and habitat census data, we recognized the crucial sites for waterbirds during wintering seasons by comparing the difference of waterbird populations at the community, foraging guild and species levels in different natural wetlands within East Dongting Lake, and then identified the crucial environmental variables affecting waterbird distributions by analyzing the relationship between waterbird populations and the environmental variables, including vegetation area, mudflat area, water area with the depth of 0-20 cm, water area with the depth of 20-50 cm, water area with the depth of 50-100 cm, water area with the depth >100 cm, growth status of vegetation (Min, Mean and Max NDVI), and the human disturbance. Results indicated that five natural wetlands, i.e., Daxiaoxi, Chunfeng, Baihu lakes, Dingzi dyke and Tanjiaweizi, were recognized as the crucial wintering sites for migratory waterbirds in the East Dongting Lake. Among the ten selected environmental variables, water areas with the depth of 0-20 cm, 20-50 cm and >100 cm, human disturbance, Min and Mean NDVIs were identified as the crucial environmental variables overall. Waterbirds at different levels exhibited significant linear relationship with certain environmental variables, with the exception of Bean goose and Lesser White-fronted goose at the species level, which showed Gaussian distribution with changes in mean NDVI. The crucial environmental variables appeared to be foraging guild- and species-specific. These findings provide significant information for managers to understand the differences of wetlands and waterbird populations within East Dongting Lake, and to make more targeted conservation efforts.

Responses to Sedimentation in Ramet Populations of the Clonal Plant Carex brevicuspis.

In aquatic ecosystems, sedimentation is an important factor that affects plant growth, mainly due to sediment depth. Clonal morphological plasticity is an effective strategy in clonal plants for acclimatization to sediment burial. To date, few studies have examined growth responses to sedimentation on the clonal plants at the ramet population level. This study aimed to explore the interactive effects of population size and burial depth on growth and clonal morphology of Carex brevicuspis. Three population sizes (2, 8, and 32 ramets) and 3 burial depths (0 cm, 5 cm, and 10 cm) were used in this experiment. Under shallow (5 cm) and deep (10 cm) burial conditions, biomass accumulation and relative growth rate (RGR) were lower than in the no burial treatment (P < 0.05). RGR of the small and medium populations was especially high compared to the large populations (P < 0.05). Biomass allocation was higher to belowground parts than aboveground parts, except for the small populations in the 5 cm burial treatments. Both shallow burial and smaller populations led to more biomass being allocated to aboveground parts. Deep burial elongated the first order spacer more than shallow burial, and sedimentation had negative effects on the second order spacer length. The number of new ramets did not decrease in the 5 or 10 cm burial treatments compared to the unburial treatment, and larger populations usually had more ramets than smaller ones; the proportion of clumping ramets was higher than the proportion of spreading ramets, and deeper burial and smaller populations led to higher proportions of spreading ramets. These results indicated that the growth of C. brevicuspis was limited by sediment burial at the ramet population level. Smaller populations enable C. brevicuspis to adjust its escape response to burial stress, may allow this species to effectively survive and widely distribute in Dongting Lake wetland.

Differential Strategies to Tolerate Flooding in Polygonum hydropiper Plants Originating From Low- and High-Elevation Habitats.

In species that occur over a wide range of flooding conditions, plant populations may have evolved divergent strategies as a consequence of long-term adaptation to local flooding conditions. In the present study, we investigated the effects of a flooding gradient on the growth and carbohydrate reserves of Polygonum hydropiper plants originating from low- and high-elevation habitats in the Dongting Lake wetlands. The results indicated that shoot length did not differ, whereas the total biomass and carbohydrate reserves were reduced under flooded compared to well-drained conditions for plants originating from both habitat types. However, shoot length, shoot mass, rhizome mass, and total biomass were lower in plants from low-elevation habitats than in those from high-elevation habitats in the flooded condition. Soluble sugar and starch contents in belowground biomass were higher in plants from low-elevation habitats than in those from high-elevation habitats independently of the water level. Therefore, P. hydropiper plants from low-elevation habitats exhibit a lower growth rate and more conservative energy strategy to cope with flooding in comparison with plants from high-elevation habitats. Differential strategies to cope with flooding among P. hydropiper populations are most likely a response to the flooding pressures of the habitat of origin and may potentially drive ecotype differentiation within species along flooding gradients.

Inhibition of litter decomposition of two emergent macrophytes by addition of aromatic plant powder.

Aromatic plants show antimicrobial activity due to their essential oils, but their effect on litter decomposition is unclear. In this study, we evaluated the biomass loss and nutrient dynamics in leaf litters of two macrophytes (Miscanthus sacchariflorus and Carex brevicuspis) with and without addition of powdered material of the aromatic plant Polygonum hydropiper or the non-aromatic plant C. brevicuspis. The two powders had similar basic chemical qualities but P. hydropiperi had a higher essential oils concentration. Leaf litters of M. sacchariflorus and C. brevicuspis were incubated with powdered P. hydropiper or C. brevicuspis (500 g m-3, 250 g m-3, and no addition) for 120 days in a mesocosm experiment. Compared with the control (no addition), P. hydropiperi addition decelerated nutrient release and litter decomposition, while C. brevicuspis addition accelerated those processes. The nitrogen concentrations in both leaf litters and the phosphorus concentration in C. brevicuspis leaf litter were increased by addition of both plant powders. The fungal biomass in both leaf litters decreased after P. hydropiperi addition, due to the antifungal activity of its essential oils. These data indicate that the aromatic plant P. hydropiperi inhibits litter decomposition via its essential oils and that such inhibition is not species-specific.

Consequences of Repeated Defoliation on Belowground Bud Banks of Carex brevicuspis (Cyperaceae) in the Dongting Lake Wetlands, China.

Despite the predominant role of bud banks in the regeneration of clonal macrophyte populations, few studies have examined the way in which clonal macrophytes adjust the demographic features of bud banks to regulate population dynamics in response to defoliation in wetlands. We investigated the density and composition of bud banks under repeated defoliation in the wetland sedge Carex brevicuspis C. B. Clarke in the Dongting Lake wetlands, China. The density and biomass of rhizome buds and shoots did not decrease significantly in response to repeated defoliation over two consecutive years. The composition of bud banks, which consisted of long and short rhizome buds, also did not change significantly in response to repeated defoliation. Nevertheless, the ramet height and the shoot, root, and rhizome mass of C. brevicuspis declined significantly under repeated defoliation. Our findings suggest that bud banks are a conservative reproductive strategy that enables C. brevicuspis to tolerate a certain amount of defoliation. The maintenance of large bud banks after repeated defoliation may enable C. brevicuspis populations to regenerate and persist in disturbed habitats. However, bud bank density of C. brevicuspis might decline in the long term because the amount of carbon stored in rhizome buds and plants is reduced by frequent defoliation.

Trade-off between allocation to reproductive ramets and rhizome buds in Carex brevicuspis populations along a small-scale elevational gradient.

The trade-off between allocation to sexual and clonal reproduction in clonal plants is influenced by a variety of environmental factors; however, it has rarely been examined under field conditions. In this study, we investigated the trade-off between two modes of reproduction in Carex brevicuspis C. B. Clarke across a small-scale elevational gradient (21-27 m a.s.l.) at the Dongting Lake wetlands, China. The proportion of biomass allocated to and the density of reproductive ramets were higher at low than at intermediate and high elevations. In contrast, the proportion of biomass allocated to and the density of rhizome buds were lower at low than at intermediate and high elevations. Redundancy analysis showed that sexual reproduction was positively correlated with soil moisture content, soil organic matter, total phosphorus, and pH, and negatively correlated with elevation and ramet density. Our findings suggested that allocation to sexual reproduction is favored in disturbed habitats with fertile soils, whereas allocation to vegetative propagation is favored in stable and competitive habitats. Trade-off between allocation to sexual reproduction and vegetative propagation along an elevational gradient might be a reproductive strategy of C. brevicuspis to adapt to the water level fluctuations in wetland habitats.

The role of seedling recruitment from juvenile populations of Carex brevicuspis (Cyperaceae) at the Dongting Lake wetlands, China.

Seedlings and vegetative ramets may contribute differentially to the recruitment of clonal populations in different growth phases, but this has rarely been investigated. In this study, we quantified the number and survivorship of seedlings and vegetative ramets monthly in juvenile and mature populations of Carex brevicuspis. During the first growing season after flooding (from October to January), 9 seedlings m(-2) (13% of all established shoots) were found in juvenile populations, while no seedlings were found in mature populations. During the second growing season before flooding (from February to May), no new seedling recruits were found either in juvenile or in mature populations. All shoots of seedlings were withered during the dormant season (January and February), but 62.5% seedlings could produce vegetative ramets in the following growing season. During the dormant season, all the early emerging ramets (sprouted in October) withered, but the later emerging ones (sprouted in November and December) survived in both mature and juvenile populations. These results indicated that seedling recruitment was only apparent in juvenile populations of C. brevicuspis. The genetic diversity in mature C. brevicuspis populations may be established in juvenile populations by seedling recruitment, and sustained in mature populations by vegetative reproduction.