Cultivated alien plants with high invasion potential are more likely to be traded online in China.

Biological invasions have become a worldwide problem, and measures to efficiently prevent and control invasions are still in development. Like many other parts of the world, China is undergoing a dramatic increase in plant invasions. Most of the currently 933 established (i.e., naturalized) plant species, of which 214 are categorized as invasive, have been introduced into China for cultivation. It is likely that many of those species are still being traded, particularly online, by plant nurseries. However, studies assessing whether naturalized and invasive species are currently being traded more or less than nonnaturalized aliens are rare. We extracted online-trade information for 13,718 cultivated alien plant taxa on 1688.com, the largest website for domestic B2B in China. We analyzed how the presence in online-nursery catalogs, the number of online nurseries that offerred the species for sale, and the product type (i.e., seeds, live plants and vegetative organs) differed among nonnaturalized, naturalized noninvasive, and invasive species. Compared to nonnaturalized taxa, naturalized noninvasive and invasive taxa were 3.7-5.2 times more likely to be available for purchase. Naturalized noninvasive and invasive taxa were more frequently offered as seeds by online nurseries, whereas nonnaturalized taxa were more frequently offered as live plants. Based on these findings, we propose that, to reduce the further spread of invasive and potentially invasive plants, implementation of plant-trade regulations and a monitoring system of the online horticultural supply chain will be essential.

The priming effects of plant leachates on dissolved organic matter degradation in water depend on leachate type and water stability.

The modification of dissolved organic matter (DOM) degradation by plant carbon inputs represents a critical biogeochemical process that controls carbon dynamics. However, the priming effects (PEs) different plant tissues induce on the degradation of DOM pools with different stabilities remain unknown. In this study, PEs, induced by different tissue leachates of Phragmites australis, were evaluated via changes in DOM components and properties of both fresh and tidal water (with different stabilities). The results showed that DOM derived from different plant tissue leachates differed in composition and bioavailability. Inputs of tissue leachates induced PEs with different intensities and directions (negative or positive) on DOM degradation of fresh and tidal water. In fresh water, the PEs of leaf and root leachates were significantly higher than those of stem and rhizome leachates. The PE direction changed for DOM degradation between fresh and tidal water. The addition of leaf and root leachates tended to induce positive PEs on DOM degradation of fresh water, while resulting in negative PEs on DOM degradation of tidal water. Negative PEs for tidal water DOM may be due to preferential utilization of microbes, high salinity, and/or the promotion of exogenous DOM production from plant tissues. The results indicate that intensity and direction of PEs induced by plant leachates depend on both leachate type and water stability. The findings highlight the necessity to examine the nature of exogenous and native DOM when interpreting the interactive processes that regulate DOM degradation.

Shifting effects of physiological integration on performance of a clonal plant during submergence and de-submergence.

BACKGROUND AND AIMS: Submergence and de-submergence are common phenomena encountered by riparian plants due to water level fluctuations, but little is known about the role of physiological integration in clonal plants (resource sharing between interconnected ramets) in their adaptation to such events. Using Alternanthera philoxeroides (alligator weed) as an example, this study tested the hypotheses that physiological integration will improve growth and photosynthetic capacity of submerged ramets during submergence and will promote their recovery following de-submergence. METHODS: Connected clones of A. philoxeroides, each consisting of two ramet systems and a stolon internode connecting them, were grown under control (both ramet systems untreated), half-submerged (one ramet system submerged and the other not submerged), fully submerged (both ramet systems submerged), half-shaded (one ramet system shaded and the other not shaded) and full-shaded (both ramet systems shaded) conditions for 30 d and then de-submerged/de-shaded for 20 d. The submerged plants were also shaded to very low light intensities, mimicking typical conditions in turbid floodwater. KEY RESULTS: After 30 d of submergence, connections between submerged and non-submerged ramets significantly increased growth and carbohydrate accumulation of the submerged ramets, but decreased the growth of the non-submerged ramets. After 20 d of de-submergence, connections did not significantly affect the growth of either de-submerged or non-submerged ramets, but de-submerged ramets had high soluble sugar concentrations, suggesting high metabolic activities. The shift from significant effects of integration on both submerged and non-submerged ramets during the submergence period to little effect during the de-submergence period was due to the quick recovery of growth and photosynthesis. The effects of physiological integration were not found to be any stronger under submergence/de-submergence than under shading/de-shading. CONCLUSIONS: The results indicate that it is not just the beneficial effects of physiological integration that are crucial to the survival of riparian clonal plants during periods of submergence, but also the ability to recover growth and photosynthesis rapidly after de-submergence, which thus allows them to spread.

Acupuncture as an adjunctive therapy for gastric ulcer: A modified Delphi consensus study.

BACKGROUND: Acupuncture is often used as an adjunctive therapy for gastric ulcer (GU). However, there is still a lack of evidence on the appropriate and optimal interventions for acupuncture. This study aimed to optimize the acupuncture treatment of gastric ulcers based on expert consensus for guiding acupuncturists in clinical practice. METHODS: To conduct this study, research evidence was gathered from databases in both Chinese and English. After discussion, preliminary clinical questions were developed. Following three rounds of multidisciplinary clinical expert consultation, the initial consensus questionnaire was formed after testing and modification by team members. A Delphi consensus was ultimately reached to answer the questionnaire and develop guidance for acupuncture treatment. A 9-point Likert-type scale was used to measure the agreement of expert consensus, where a score of 80% between 7 and 9 was defined as "agreement." RESULTS: After two rounds of Delphi voting, a total of 35 items reached an agreement. These items can be roughly divided into 6 domains. According to expert consensus, the application of acupuncture for gastric ulcer should follow a semistandardized approach. Based on the syndrome differentiation, the main acupoints recommended are Zusanli (ST36), Zhongwan (CV12), Neiguan (PC6), and Sanyinjiao (SP6), while the adjunct acupoints include Taichong (LR3), Guanyuan (CV4), Xuehai (SP10), and Taixi (KI3). In the experience of experts, adverse events associated with acupuncture are typically mild and often manifest as subcutaneous hematomas. CONCLUSION: There is a lack of definitive acupuncture guidelines that can effectively determine the optimal therapeutic approach for the treatment of gastric ulcer. This expert consensus provides recommendations for clinical research and practice of acupuncture, with a particular focus on the selection of acupoints. However, further exploration through rigorous studies is necessary due to the limited availability of clinical evidence.

Transgenerational Herbivory Effects on Performance of Clonal Offspring of the Invasive Plant Alternanthera philoxeroides.

Interactions between alien plants and local enemies in introduced ranges may determine plant invasion success. However, little is known about whether herbivory-induced responses are transmitted across vegetative generations of plants and whether epigenetic changes are involved during this process. In a greenhouse experiment, we examined the effects of herbivory by the generalist herbivore Spodoptera litura on the growth, physiology, biomass allocation and DNA methylation level of the invasive plant Alternanthera philoxeroides in the first- (G1), second- (G2) and third-generation (G3). We also tested the effects of root fragments with different branching orders (i.e., the primary- or secondary-root fragments of taproots) of G1 on offspring performance. Our results showed that G1 herbivory promoted the growth of the plants in G2 that sprouted from the secondary-root fragments of G1 but had a neutral or negative effect on the growth of the plants in G2 from the primary-root fragments. The growth of plants in G3 was significantly reduced by G3 herbivory but not affected by G1 herbivory. Plants in G1 exhibited a higher level of DNA methylation when they were damaged by herbivores than when they were not, while neither plants in G2 nor G3 showed herbivory-induced changes in DNA methylation. Overall, the herbivory-induced growth response within one vegetative generation may represent the rapid acclimatization of A. philoxeroides to the unpredictable generalist herbivores in the introduced ranges. Herbivory-induced trans-generational effects may be transient for clonal offspring of A. philoxeroides, which can be influenced by the branching order of taproots, but be less characterized by DNA methylation.

Acclimatory responses of Arabidopsis to fluctuating light environment: comparison of different sunfleck regimes and accessions.

Acclimation to fluctuating light environment with short (lasting 20 s, at 650 or 1,250 µmol photons m(-2) s(-1), every 6 or 12 min) or long (for 40 min at 650 µmol photons m(-2) s(-1), once a day at midday) sunflecks was studied in Arabidopsis thaliana. The sunfleck treatments were applied in the background daytime light intensity of 50 µmol photons m(-2) s(-1). In order to distinguish the effects of sunflecks from those of increased daily irradiance, constant light treatments at 85 and 120 µmol photons m(-2) s(-1), which gave the same photosynthetically active radiation (PAR) per day as the different sunfleck treatments, were also included in the experiments. The increased daily total PAR in the two higher constant light treatments enhanced photosystem II electron transport and starch accumulation in mature leaves and promoted expansion of young leaves in Columbia-0 plants during the 7-day treatments. Compared to the plants remaining under 50 µmol photons m(-2) s(-1), application of long sunflecks caused upregulation of electron transport without affecting carbon gain in the form of starch accumulation and leaf growth or the capacity of non-photochemical quenching (NPQ). Mature leaves showed marked enhancement of the NPQ capacity under the conditions with short sunflecks, which preceded recovery and upregulation of electron transport, demonstrating the initial priority of photoprotection. The distinct acclimatory responses to constant PAR, long sunflecks, and different combinations of short sunflecks are consistent with acclimatory adjustment of the processes in photoprotection and carbon gain, depending on the duration, frequency, and intensity of light fluctuations. While the responses of leaf expansion to short sunflecks differed among the seven Arabidopsis accessions examined, all plants showed NPQ upregulation, suggesting limited ability of this species to utilize short sunflecks. The increase in the NPQ capacity was accompanied by reduced chlorophyll contents, higher levels of the xanthophyll-cycle pigments, faster light-induced de-epoxidation of violaxanthin to zeaxanthin and antheraxanthin, increased amounts of PsbS protein, as well as enhanced activity of superoxide dismutase. These acclimatory mechanisms, involving reorganization of pigment-protein complexes and upregulation of other photoprotective reactions, are probably essential for Arabidopsis plants to cope with photo-oxidative stress induced by short sunflecks without suffering from severe photoinhibition and lipid peroxidation.

Nutrient Inputs Alleviate Negative Effects of Early and Subsequent Flooding on Growth of Polygonum hydropiper With the Aid of Adventitious Roots.

Riparian plants are exposed to harmful stress induced by flooding, which is often accompanied by eutrophication in the Three Gorges Reservoir Region. The phenomenon is mainly caused by domestic sewage discharges, slow water flow, and agricultural fertilizer pollution. Simulating abiotic stress, such as flooding at the initial period, can act as a signal and induce positive responses of plants to subsequent severe stress. In addition, eutrophication supplies nutrients, provides a favorable environment in the early stages of plant, and facilitates good performance in later development. However, whether early flooding (with or without eutrophication) acts as positive cue or as stress on plants at different developmental stages remains unclear. To address this question, seeds of Polygonum hydropiper were collected from low and high elevations in the hydro-fluctuation belt of the Three Gorges Reservoir Region. Plants germinated from these seeds were subjected to shallower and shorter early flooding treatments with or without eutrophication. Subsequently, plants were subjected to deeper and longer flooding treatments with or without eutrophication. Early flooding and eutrophic flooding significantly induced generation of adventitious roots, suggesting morphological adaptation to flooding. Although early flooding and eutrophic flooding treatments did not increase plant biomass in subsequent treatments compared with control, stem length, length and width of the 1st fully expanded leaf, and biomass of plants in the early eutrophic treatment were higher than these of the early flooding treatment plants. These results suggest a negative lag-effect of early flooding, and also indicate that nutrient inputs can alleviate such effects. Similarly, subsequent eutrophic flooding also enhanced plant growth compared with subsequent flooding, showing significantly higher values of leaf traits and adventitious root number. Plants originated from low elevation had significantly higher functional leaf length and stem biomass compared with those from high elevation. These results suggest that nutrient inputs can alleviate negative effects of early and subsequent flooding on growth of P. hydropiper with the generation of adventitious roots.

Roles of clonal parental effects in regulating interspecific competition between two floating plants.

Parental effects can influence offspring fitness, which may further impact interspecific competition. However, few studies have tested the role of clonal parental effects in regulating interspecific interactions and examined the underlying mechanisms. We conducted two consecutive experiments with two clonal plants (Pistia stratiotes and Eichhornia crassipes). In the first experiment, the mother ramet of P. stratiotes and E. crassipes were grown in two nutrient levels and treated with a DNA demethylation reagent (5-azacytidine) or not. In the second experiment, the offspring ramets from each of the four treatments in the first experiment were grown alone (no competition) or with a heterospecific neighbor (with interspecific competition). We found no parental nutrient effect on the competitive ability of E. crassipes, but a significant parental nutrient effect of both E. crassipes and P. stratiotes on the competitive ability of P. stratiotes. Furthermore, the parental nutrient effect of P. stratiotes on the competitive ability of P. stratiotes varied depending on the DNA methylation status of both P. stratiotes and E. crassipes. These clonal parental effects were related to resource provisioning and/or DNA methylation. We conclude that clonal parental nutrient effects can regulate interspecific competition between P. stratiotes and E. crassipes by altering the competitive ability of P. stratiotes. Both resource provisioning and epigenetic mechanisms can be involved in these clonal parental effects. By regulating interspecific competition, clonal parental effects may further influence species coexistence, community structure, and ecosystem functioning.

Parental effects driven by resource provisioning in Alternanthera philoxeroides-A simulation case study.

Parental environmental effects can be a rapid and effective means for clonal plants in response to temporally or spatially varying environments. However, few studies have quantitatively measured the ecological significance of parental effects in aquatic clonal plants. In this study, we developed a two-generation (parent-offspring) growth model to examine the parental effects of nitrogen (N) conditions on summed and mean performance of clonal offspring of one wetland species Alternanthera philoxeroides. We also examined the role of survival status and developmental stage of clonal offspring in the consequence of parental effects in aquatic clonal plants. Our results indicated direct evidence that (1) there were significant non-linear correlations between the performance of parental plants and initial status of clonal offspring (i.e., the mass and number of clonal propagules); (2) parental N effects on the summed performance of clonal offspring were content-dependent (i.e., there were significant interactions between parental and offspring N effects), while parental effects on the mean performance of offspring were independent of offspring conditions; (3) parental effects mainly occurred at the early development stage of clonal offspring, and then gradually declined at the late stage; (4) the context-dependent parental effects on the summed performance of clonal offspring gradually strengthened when offspring survival was high. The mathematical models derived from the experimental data may help researchers to not only deeply explore the ecological significance of parental environmental effects in aquatic clonal plants, but also to reveal the importance of potential factors that have been often neglected in empirical studies.

Recovery dynamics of growth, photosynthesis and carbohydrate accumulation after de-submergence: a comparison between two wetland plants showing escape and quiescence strategies.

BACKGROUND AND AIMS: The capacity for fast-growth recovery after de-submergence is important for establishment of riparian species in a water-level-fluctuation zone. Recovery patterns of two wetland plants, Alternanthera philoxeroides and Hemarthria altissima, showing 'escape' and 'quiescence' responses, respectively, during submergence were investigated. METHODS: Leaf and root growth and photosynthesis were monitored continuously during 10 d of recovery following 20 d of complete submergence. Above- and below-ground dry weights, as well as carbohydrate concentrations, were measured several times during the experiment. KEY RESULTS: Both species remobilized stored carbohydrate during submergence. Although enhanced internode elongation depleted the carbohydrate storage in A. philoxeroides during submergence, this species resumed leaf growth 3 d after de-submergence concomitant with restoration of the maximal photosynthetic capacity. In contrast, some sucrose was conserved in shoots of H. altissima during submergence, which promoted rapid re-growth of leaves 2 d after de-submergence and earlier than the full recovery of photosynthesis. The recovery of root growth was delayed by 1-2 d compared with leaves in both species. CONCLUSIONS: Submergence tolerance of the escape and quiescence strategies entails not only the corresponding regulation of growth, carbohydrate catabolism and energy metabolism during submergence but also co-ordinated recovery of photosynthesis, growth and carbohydrate partitioning following de-submergence.

Effects of salinity and concomitant species on growth of Phragmites australis populations at different levels of genetic diversity.

In plant communities, genetic diversity among dominant species can not only affect the fitness of the population, but also interactions with concomitant species. Soil salinity is a common factor that influences plant growth in estuarine wetlands. However, few studies have tested whether their high genetic diversity will be beneficial for the resistance of plant populations to salinity and the presence of concomitant plants. Four different genotypes of Phragmites australis, a dominant species of the Yellow River Delta in China, were selected to construct populations with three different genotypic levels. These populations were planted either with or without concomitant species and were subjected to control or salinity treatments. At the end of treatments, growth variables of P. australis populations were measured. In response to soil salinity, the total biomass of 1-, 2-, and 4-genotype populations decreased by 35%, 24%, and 13%, respectively, indicating higher resistance of P. australis populations with high genetic diversity. Correspondingly, 2-, and 4-genotype populations showed higher biomass allocation to roots, which can maintain adequate water uptake for plants. The biomass accumulation of 1-genotype populations with concomitant plants was significantly lower compared with populations without concomitant plants; however, no significant difference was found for 4-genotype populations between both control and salinity treatments, suggesting their higher capacities when coexisting with concomitant species. However, the genotypic level of populations did not significantly affect their biomass accumulation. High genetic diversity is greatly beneficial for the resistance of P. australis populations to salinity and coexistence with other plants. This information should be considered in the construction or restoration of this species in estuarine wetlands.

Photosynthetic acclimation is important for post-submergence recovery of photosynthesis and growth in two riparian species.

BACKGROUND AND AIMS: Concomitant increases in O(2) and irradiance upon de-submergence can cause photoinhibition and photo-oxidative damage to the photosynthetic apparatus of plants. As energy and carbohydrate supply from photosynthesis is needed for growth, it was hypothesized that post-submergence growth recovery may require efficient photosynthetic acclimation to increased O(2) and irradiance to minimize photo-oxidative damage. The hypothesis was tested in two flood-tolerant species: a C(3) herb, Alternanthera philoxeroides; and a C(4) grass, Hemarthria altissima. The impact of low O(2) and low light, typical conditions in turbid floodwater, on post-submergence recovery was assessed by different flooding treatments combined with shading. METHODS: Experiments were conducted during 30 d of flooding (waterlogging or submergence) with or without shading and subsequent recovery of 20 d under growth conditions. Changes in dry mass, number of branches/tillers, and length of the longest internodes and main stems were recorded to characterize growth responses. Photosynthetic parameters (photosystem II efficiency and non-photochemical quenching) were determined in mature leaves based on chlorophyll a fluorescence measurements. KEY RESULTS: In both species growth and photosynthesis recovered after the end of the submergence treatment, with recovery of photosynthesis (starting shortly after de-submergence) preceding recovery of growth (pronounced on days 40-50). The effective quantum yield of photosystem II and non-photochemical quenching were diminished during submergence but rapidly increased upon de-submergence. Similar changes were found in all shaded plants, with or without flooding. Submerged plants did not suffer from photoinhibition throughout the recovery period although their growth recovery was retarded. CONCLUSIONS: After sudden de-submergence the C(3) plant A. philoxeroides and the C(4) plant H. altissima were both able to maintain the functionality of the photosynthetic apparatus through rapid acclimation to changing O(2) and light conditions. The ability for photosynthetic acclimation may be essential for adaptation to wetland habitats in which water levels fluctuate.

Growth responses of eight wetland species to water level fluctuation with different ranges and frequencies.

Fluctuation range and frequency are two important components of water level fluctuation, but their effects on wetland plants have not been evaluated separately. We subjected eight wetland species to a control treatment with static water level and fluctuation treatments with different ranges or frequencies to examine their effects on plant growth. Acorus calamus, Butomus umbellatus and Iris wilsonii showed high survival rates in all treatments with various fluctuation ranges and frequencies. Their survival rates were higher at the medium fluctuation frequency than at the low and high frequencies, suggesting beneficial effects of the medium frequency. In the experiment comparing the fluctuation ranges, A. calamus and I. wilsonii could maintain the capacity for asexual propagation and accumulate higher biomass compared with the control plants, while biomass of the other six species dramatically decreased. In the experiment comparing fluctuation frequency, species with relatively high survival rates (≥ 50%) maintained or increased the capacity of asexual propagation, and A. calamus and I. wilsonii allocated relatively more biomass to roots, which may enhance plant growth and survival. In contrast, these species did not show increased biomass allocation to shoots in response to both fluctuation range and frequency, presumably because shoots are prone to mechanical damage caused by streaming floodwater. Taken together, biomass accumulation in roots rather than in shoots and the ability to asexually propagate are important for the survival of these species during water fluctuation.

Astragaloside suppresses apoptosis of the podocytes in rats with diabetic nephropathy via miR-378/TRAF5 signaling pathway.

AIMS: Apoptosis of podocytes plays a crucial role in diabetic nephropathy (DN) development, and astragaloside (AS-IV) has a significant impact on podocyte apoptosis. This study aims to explore the effect of AS-IV on diabetic nephropathy progression. MATERIALS AND METHODS: The diabetic nephropathy model was established in rats with streptozotocin (STZ) injection. The albuminuria was examined by using the enzyme linked immunosorbent assay (ELISA). The expression of miR-378, tumor-necrosis factor (TNF) receptor (TNFR)-associated factor 5 (TRAF5) mRNA and protein was analyzed by qRT-PCR and western blot, respectively. Cell transfection was conducted for modulating endogenous expression of miR-378. Dual luciferase reporter assay was used to evaluate the interaction between miR-378 and TRAF5. The terminal deoxynucleotidy transferase dUTP nick end labeling (TUNEL) staining assay was performed for apoptosis detection. KEY FINDINGS: AS-IV protected diabetic rats from developing into diabetic nephropathy. The expression of miR-378 was down-regulated in diabetic nephropathy rats, which was reversed by AS-IV. AS-IV enhanced the expression of miR-378 in podocytes treated with high glucose. MiR-378 negatively regulated TRAF5. AS-IV inhibited the expression of TRAF5 through miR-378. AS-IV suppressed apoptosis of podocytes via targeting miR-378. SIGNIFICANCE: AS-IV suppresses apoptosis of the podocytes through the miR-378/TRAF5 signaling pathway, and thereby repressing diabetic nephropathy development.

Herbivory-induced maternal effects on growth and defense traits in the clonal species Alternanthera philoxeroides.

Plants have evolved a variety of defense traits against foliar herbivory, including the production of primary and secondary metabolites, the allocation of chemical compounds, and morphological plasticity. Using two vegetative generations of the invasive clonal species Alternanthera philoxeroides, we investigated the effects of maternal and offspring herbivory by Planococcus minor on the integrative defense strategy of plants. Herbivory severely inhibited leaf, stolon and root growth, as well as the production of primary metabolites (soluble sugars, starch, and total non-structural carbohydrates in stolons), and decreased average leaf area and specific leaf area of the second-generation A. philoxeroides. The changes in growth measures of the first-generation A. philoxeroides with herbivory were consistent with that of the second generation. By contrast, herbivory basically did not affect the concentration of non-structural carbohydrate compounds in the roots, and even increased the concentrations of N and total phenols in taproots. Furthermore, herbivory-induced maternal effects also reduced the growth of the second-generation plants. The results suggest that A. philoxeroides is capable of adapting to herbivory by P. minor, mainly via the allocation of available resources in belowground organs, and that the herbivory effect can persist across vegetative generations. These features may potentially improve the regeneration and tolerance of A. philoxeroides after a short-term herbivory.

Heterogeneous water supply affects growth and benefits of clonal integration between co-existing invasive and native Hydrocotyle species.

Spatial patchiness and temporal variability in water availability are common in nature under global climate change, which can remarkably influence adaptive responses of clonal plants, i.e. clonal integration (translocating resources between connected ramets). However, little is known about the effects of spatial patchiness and temporal heterogeneity in water on growth and clonal integration between congeneric invasive and native Hydrocotyle species. In a greenhouse experiment, we subjected severed or no severed (intact) fragments of Hydrocotyle vulgaris, a highly invasive species in China, and its co-existing, native congener H. sibthorpioides to different spatial patchiness (homogeneous and patchy) and temporal interval (low and high interval) in water supply. Clonal integration had significant positive effects on growth of both species. In the homogeneous water conditions, clonal integration greatly improved the growth in fragments of both species under low interval in water. However, in the patchy water conditions, clonal integration significantly increased growth in both ramets and fragments of H. vulgaris under high interval in water. Therefore, spatial patchiness and temporal interval in water altered the effects of clonal integration of both species, especially for H. vulgaris. The adaptation of H. vulgaris might lead to invasive growth and potential spread under the global water variability.

Do Amplitudes of Water Level Fluctuations Affect the Growth and Community Structure of Submerged Macrophytes?

Submerged macrophytes are subjected to potential mechanical stresses associated with fluctuating water levels in natural conditions. However, few experimental studies have been conducted to further understand the effects of water level fluctuating amplitude on submerged macrophyte species and their assemblages or communities. We designed a controlled experiment to investigate the responses of three submerged macrophyte species (Hydrilla verticillata, Ceratophyllum demersum and Elodea nuttallii) and their combinations in communities to three amplitudes (static, ± 30 cm, ± 60 cm) of water level fluctuations. Results showed that water level fluctuating amplitude had little effects on the community performance and the three tested species responded differently. H. verticillata exhibited more growth in static water and it was negatively affected by either of the water level fluctuations amplitude, however, growth parameters of H. verticillata in two fluctuating water level treatments (i.e., ± 30 cm, ± 60 cm) were not significantly different. On the other hand, the growth of C. demersum was not significantly correlated with different amplitude treatments. However, it became more abundant when water levels fluctuated. E. nuttallii was inhibited by the two fluctuating water level treatments, and was less in growth parameters compared to the other species especially in water level fluctuating conditions. The inherent differences in the adaptive capabilities of the tested species indicate that C. demersum or other species with similar responses may be dominant species to restore submerged macrophyte communities with great fluctuating water levels. Otherwise, H. verticillata, E. nuttallii or other species with similar responses could be considered for constructing the community in static water conditions.

Effects of Spatial Patch Arrangement and Scale of Covarying Resources on Growth and Intraspecific Competition of a Clonal Plant.

Spatial heterogeneity in two co-variable resources such as light and water availability is common and can affect the growth of clonal plants. Several studies have tested effects of spatial heterogeneity in the supply of a single resource on competitive interactions of plants, but none has examined those of heterogeneous distribution of two co-variable resources. In a greenhouse experiment, we grew one (without intraspecific competition) or nine isolated ramets (with competition) of a rhizomatous herb Iris japonica under a homogeneous environment and four heterogeneous environments differing in patch arrangement (reciprocal and parallel patchiness of light and soil water) and patch scale (large and small patches of light and water). Intraspecific competition significantly decreased the growth of I. japonica, but at the whole container level there were no significant interaction effects of competition by spatial heterogeneity or significant effect of heterogeneity on competitive intensity. Irrespective of competition, the growth of I. japonica in the high and the low water patches did not differ significantly in the homogeneous treatments, but it was significantly larger in the high than in the low water patches in the heterogeneous treatments with large patches. For the heterogeneous treatments with small patches, the growth of I. japonica was significantly larger in the high than in the low water patches in the presence of competition, but such an effect was not significant in the absence of competition. Furthermore, patch arrangement and patch scale significantly affected competitive intensity at the patch level. Therefore, spatial heterogeneity in light and water supply can alter intraspecific competition at the patch level and such effects depend on patch arrangement and patch scale.

Effects of patch contrast and arrangement on benefits of clonal integration in a rhizomatous clonal plant.

The availabilities of light and soil water resources usually spatially co-vary in natural habitats, and the spatial pattern of such co-variation may affect the benefits of physiological integration between connected ramets of clonal plants. In a greenhouse experiment, we grew connected or disconnected ramet pairs [consisting of a proximal (relatively old) and a distal (relative young) ramet] of a rhizomatous herb Iris japonica in four heterogeneous environments differing in patch arrangement (reciprocal vs. parallel patchiness of light and soil water) and patch contrast (high vs. low contrast of light and water). Biomass of the proximal part, distal part and clonal fragment of I. japonica were all significantly greater in the intact than in the severed treatment, in the parallel than in the reciprocal patchiness treatment and in the high than in the low contrast treatment, but the effect of severing the connection between ramet pairs did not depend on patch arrangement or contrast. Severing the connection decreased number of ramets of the distal part and the clonal fragment in the parallel patchiness arrangement, but not in the reciprocal patchiness arrangement. Therefore, the spatial arrangement of resource patches can alter the effects of clonal integration on asexual reproduction in I. japonica.

[Effects of light intensity contrast on clonal integration of Spartina anglica].

We conducted a greenhouse experiment to test how clonal integration affected the growth responses of Spartina anglica to light intensity heterogeneity in light availability and whether such effects depended on contrast light intensity. The experiment had three homogeneous treatments and two heterogeneous treatments. In the homogeneous treatments, both ramets of a connected ramet pair were unshaded (high light intensity), moderately shaded (medium light intensity, 70% of the high light intensity) and deeply shaded (low light intensity, 30% of the high light intensity). In the heterogeneous treatments, one ramet of a pair was unshaded, but its connected ramet was either moderately shaded (low light intensity contrast) or deeply shaded (high light intensity contrast). In the homogeneous treatments, biomass of S. anglica was significantly higher in the high light intensity treatment than in the medium and low light intensity treatments. Number of leaves, root length, and total biomass were significantly higher in the shaded ramet in the heterogeneous treatment with low light intensity contrast than in the ramet in the homogeneous treatment with medium light intensity. Final size and mass were not significantly different between the unshaded ramet in the heterogeneous treatment with low light intensity contrast and the ramets in the homogeneous high light intensity treatment. These results suggested that clonal integration benefitted a shaded ramet with little cost to an unshaded ramet when contrast in light intensity was low. However, effects of clonal integration were not significant when contrast was high. It therefore appeared that effects of clonal integration on the growth of S. anglica did not increase with increasing light intensity contrast. In natural habitats, clonal integration might improve growth of S. anglica when its ramets are moderately shaded by other plants but not when they are deeply shaded.