Clonal integration of stress signal induces morphological and physiological response of root within clonal network.

Clonal integration of defense or stress signal induced systemic resistance in leaf of interconnected ramets. However, similar effects of stress signal in root are poorly understood within clonal network. Clonal fragments of Centella asiaticas with first-young, second-mature, third-old and fourth-oldest ramets were used to investigate transportation or sharing of stress signal among interconnected ramets suffering from low water availability. Compared with control, oxidative stress in root of the first-young, second-mature and third-old ramets was significantly alleviated by exogenous ABA application to the fourth-oldest ramets as well as enhancement of antioxidant enzyme (SOD, POD, CAT and APX) activities and osmoregulation ability. Surface area and volume in root of the first-young ramets were significantly increased and total length in root of the third-old ramets was significantly decreased. POD activity in root of the fourth-oldest and third-old ramets was significantly enhanced by exogenous ABA application to the first-young ramets. Meanwhile, total length and surface area in root of the fourth-oldest and third-old ramets were significantly decreased. Ratio of belowground to aboveground biomass in the whole clonal fragments was significantly increased by exogenous ABA application to the fourth-oldest or first-young ramets. It is suggested that transportation or sharing of stress signal may induce systemic resistance in root of interconnected ramets. Specially, transportation or sharing of stress signal against phloem flow was observed in the experiment. Possible explanation is that rapid recovery of foliar photosynthesis in first-young ramets subjected to exogenous ABA application can partially reverse phloem flow within clonal network. Thus, our experiment provides insight into ecological implication on clonal integration of stress signal.

Soil heterogeneity in the horizontal distribution of microplastics influences productivity and species composition of plant communities.

Contamination of soils by microplastics can have profound ecological impacts on terrestrial ecosystems and has received increasing attention. However, few studies have considered the impacts of soil microplastics on plant communities and none has tested the impacts of spatial heterogeneity in the horizontal distribution of microplastics in the soil on plant communities. We grew experimental plant communities in soils with either a homogeneous or a heterogeneous distribution of each of six common microplastics, i.e., polystyrene foam (EPS), polyethylene fiber (PET), polyethylene bead (HDPE), polypropylene fiber (PP), polylactic bead (PLA) and polyamide bead (PA6). The heterogeneous treatment consisted of two soil patches without microplastics and two with a higher (0.2%) concentration of microplastics, and the homogeneous treatment consisted of four patches all with a lower (0.1%) concentration of microplastics. Thus, the total amounts of microplastics in the soils were exactly the same in the two treatments. Total and root biomass of the plant communities were significantly higher in the homogeneous than in the heterogeneous treatment when the microplastic was PET and PP, smaller when it was PLA, but not different when it was EPS, HDPE or PA6. In the heterogeneous treatment, total and root biomass were significantly smaller in the patches with than without microplastics when the microplastic was EPS, but greater when the microplastic was PET or PP. Additionally, in the heterogeneous treatment, root biomass was significantly smaller in the patches with than without microplastics when the microplastic was HDPE, and shoot biomass was also significantly smaller when the microplastic was EPS or PET. The heterogeneous distribution of EPS in the soil significantly decreased community evenness, but the heterogeneous distribution of PET increased it. We conclude that soil heterogeneity in the horizontal distribution of microplastics can influence productivity and species composition of plant communities, but such an effect varies depending on microplastic chemical composition (types) and morphology (shapes).

Clonal integration benefits an invader in heterogeneous environments with reciprocal patchiness of resources, but not its native congener.

Many of the world's most invasive plants are clonal, and clonal functional traits are suggested to contribute to their invasiveness. Clonal integration is one of the most important clonal functional traits, but it is still unclear whether clonal integration can benefit invasive alien clonal plants more than native ones in heterogeneous environments with reciprocal patchiness of resources and whether invasive plants show a higher capacity of division of labor than native ones in such environments. We grew connected (allowing clonal integration) and disconnected (preventing clonal integration) ramet pairs of an invasive plant Wedelia trilobata and its occurring native congener W. chinensis in the environment consisting of reciprocal patches of light and soil nutrients (i.e., a high-light but low-nutrient patch and a low-light but high-nutrient patch). Clonal integration greatly promoted the growth of the invasive species, but had no significant effect on the native one. Both invasive and native species showed division of labor in terms of morphology, biomass allocation, and/or photosynthetic physiology, but the capacity of labor division did not differ between the invasive and the native species. We conclude that in heterogeneous environments consisting of reciprocal patches of resources, which are common in nature, clonal integration can confer invasive plants a competitive advantage over natives, but this difference is not related to their capacity of labor division. This study highlights the importance of clonal integration for plants in heterogeneous environments and suggests that clonal integration can contribute to the invasion success of alien clonal plants.

Effects of nutrient supply on leaf stoichiometry and relative growth rate of three stoloniferous alien plants.

Different nutrient supply brings about changes in leaf stoichiometry, which may affect growth rate and primary production of plants. Invasion of alien plants is a severe threat to biodiversity and ecosystem worldwide. A pot experiment was conducted by using three stoloniferous alien plants Wedelia trilobata, Alternanther philoxeroides and Hydrocotyle vulgaris to investigate effects of nutrient supply on their leaf stoichiometry and relative growth rate. Different nitrogen or phosphorus supply was applied in the experiment (N1:1 mmol L-1, N2:4 mmol L-1, and N3:8 mmol L-1, P1:0.15 mmol L-1, P2:0.6 mmol L-1 and P3:1.2 mmol L-1). Nitrogen and phosphorus concentrations in leaves of the three alien plants significantly increased with increase of nitrogen supply. With increase of phosphorus supply, nitrogen or phosphorus concentration of leaf was complex among the three alien plants. N:P ratio in leaf of the three alien plants subjected to different levels of nutrient supply was various. A positive correlation between relative growth rate and N:P ratio of the leaf is observed in W. trilobata and A. philoxeroides suffering from N-limitation. A similar pattern was not observed in Hydrocotyle vulgaris. We tentatively concluded that correlations between relative growth rate and N: P ratio of the leaf could be affected by species as well as nutrient supply. It is suggested that human activities, invasive history, local abundance of species et al maybe play an important role in the invasion of alien plants as well as relative growth rate.

Distinct responses of frond and root to increasing nutrient availability in a floating clonal plant.

Current knowledge on responses of aquatic clonal plants to resource availability is largely based on studies manipulating limited resource levels, which may have failed to capture the "big picture" for aquatic clonal plants in response to resource availability. In a greenhouse experiment, we grew the floating clonal plant Spirodela polyrhiza under ten nutrient levels (i.e., 1/64×, 1/32×, 1/16×, 1/8×, 1/4×, 1/2×, 1×, 2×, 4× and 8×full-strength Hoagland solution) and examined their responses in terms of clonal growth, morphology and biomass allocations. The responses of total biomass and number of ramets to nutrient availability were unimodal. A similar pattern was found for frond mass, frond length and frond width, even though area per frond and specific frond area fluctuated greatly in response to nutrient availability. In contrast, the responses of root mass and root length to nutrient availability were U-shaped. Moreover, S. polyrhiza invested more to roots under lower nutrient concentrations. These results suggest that nutrient availability may have distinct influences on roots and fronds of the aquatic clonal plant S. polyrhiza, resulting in a great influence on the whole S. polyrhiza population.

Growth and Morphological Responses of Duckweed to Clonal Fragmentation, Nutrient Availability, and Population Density.

Connected ramets of aquatic clonal plants are susceptible to fragmentation by disturbance. Such clonal fragmentation may interact with nutrient availability and individual density to affect growth and morphology of aquatic clonal plants. We grew the widespread floating clonal plant Spirodela polyrhiza (duckweed) under three levels of population density (low, medium, or high), two levels of nutrient availability (low or high), and two levels of clonal fragmentation (with or without). Clonal fragmentation and high nutrients increased biomass and ramet number, but decreased frond width, frond length, and specific frond area of S. polyrhiza. Increasing population density decreased growth (biomass and ramet number) and frond and root size, and increased frond thickness of individual ramets of S. polyrhiza. The negative effect of population density on growth of S. polyrhiza was greater under high than under low nutrient availability. Furthermore, the negative effect of population density on total mass and frond mass of S. polyrhiza was greater with fragmentation than without. These results suggest that clonal fragmentation, nutrient availability and population density can interact to affect growth and morphology of clonal floating plants. Competition for nutrients and space, rather than light, may be the mechanisms underlying reduced growth of clonal floating plants. As clonal fragmentation can increase biomass and ramet production of S. polyrhiza, disturbance that potentially causes clonal fragmentation cannot be recommended as a measure to limit the spread of clonal floating plants.

Clonal integration ameliorates the carbon accumulation capacity of a stoloniferous herb, Glechoma longituba, growing in heterogenous light conditions by facilitating nitrogen assimilation in the rhizosphere.

BACKGROUND AND AIMS: Enhanced availability of photosynthates increases nitrogen (N) mineralization and nitrification in the rhizosphere via rhizodeposition from plant roots. Under heterogeneous light conditions, photosynthates supplied by exposed ramets may promote N assimilation in the rhizosphere of shaded, connected ramets. This study was conducted to test this hypothesis. METHODS: Clonal fragments of the stoloniferous herb Glechoma longituba with two successive ramets were selected. Mother ramets were subjected to full sunlight and offspring ramets were subjected to 80 % shading, and the stolon between the two successive ramets was either severed or left intact. Measurements were taken of photosynthetic and growth parameters. The turnover of available soil N was determined together with the compostion of the rhizosphere microbial community. KEY RESULTS: The microbial community composition in the rhizosphere of shaded offspring ramets was significantly altered by clonal integration. Positive effects of clonal integration were observed on NAGase activity, net soil N mineralization rate and net soil N nitrification rate. Increased leaf N and chlorophyll content as well as leaf N allocation to the photosynthetic machinery improved the photosynthetic capability of shaded offspring ramets when the stolon was left intact. Clonal integration improved the growth performance of shaded, connected offspring ramets and whole clonal fragments without any cost to the exposed mother ramets. CONCLUSIONS: Considerable differences in microbial community composition caused by clonal integration may facilitate N assimilation in the rhizosphere of shaded offspring ramets. Increased N content in the photosynthetic machinery may allow pre-acclimation to high light conditions for shaded offspring ramets, thus promoting opportunistic light capture. In accordance with the theory of the division of labour, it is suggested that clonal integration may ameliorate the carbon assimilation capacity of clonal plants, thus improving their fitness in temporally and spatially heterogeneous habitats.