Clonal integration benefits invasive alien plants under water variability in a native community

Aims Many invasive alien plant species are clonal and can greatly propagate and spread through clonal integration (sharing resources between connected ramets) in heterogeneous and variable environments. Here, we tested whether water variability influences clonal integration of invasive alien plant species and consequently facilitates their growth and dominance in a native community.     

The invasive plant Alternanthera philoxeroides benefits from clonal integration in response to defoliation

Many notorious alien invasive plants have the capacity for vigorous clonal growth, and clonal integration may contribute to their invasiveness in response to various disturbances. Here, it is hypothesized that clonal integration affects the growth, biomass allocation, physiology, and compensatory response of the alien invasive clonal plant Alternanthera philoxeroides when faced with defoliation. To test these hypotheses, a growth experiment was conducted to investigate the effect of clonal integration on the responses of A. philoxeroides to different levels of defoliation. Daughter ramets that had been grown with stolon connections that were either severed from or connected to the mother plant were subjected to four defoliation levels: 0 (control), 30% (mild), 60% (moderate) and 90% (heavy) removal of leaf tissue. Defoliation greatly decreased growth (total biomass, number of ramets and total stolon length) but increased the maximum quantum yield of photosystem II (F_v/F_m) of daughter ramets. Clonal integration significantly increased growth, F_v/F_m and contents of non-structural carbohydrates (soluble sugars and total non-structural carbohydrates) of A. philoxeroides, and these effects were larger under heavier defoliation. Moreover, clonal integration markedly reduced the shoot/root ratio of A. philoxeroides, and these effects tended to increase with increasing levels of defoliation. These results support our hypothesis that A. philoxeroides benefits from clonal integration in response to defoliation, suggesting that clonal integration may be closely related to the invasiveness of A. philoxeroides in natural habitats with frequent disturbances.     

Differences in physiological integration between invasive and noninvasive introduced clonal species of Carpobrotus

AimsClonal growth is associated with invasiveness in introduced plant species, but few studies have compared invasive and noninvasive introduced clonal species to investigate which clonal traits may underlie invasiveness. To test the hypothesis that greater capacity to increase clonal growthviaphysiological integration of connected ramets increases invasiveness in clonal plants, we compared the effects of severing connections on accumulation of mass in the two species of the creeping, succulent, perennial, herbaceous genusCarpobrotusthat have been introduced on sand dunes along the Pacific Coast of northern California, the highly invasive speciesCarpobrotus edulisand the co-occurring, noninvasive speciesCarpobrotus chilensis.     

芦苇克隆整合对石油污染湿地土壤微生物群落结构和生物量的影响

克隆植物形体相连的无性个体(分株)之间可以进行水分、养分和光合产物的传递和共享,并且这种克隆整合可以显著提高分株对环境胁迫的耐受能力,从而可能进一步影响分株周围的土壤微生物群落结构和生物量。尽管国内外已经开展了大量有关克隆整合对分株耐受胁迫能力影响的研究,但克隆整合对土壤微生物群落结构和生物量影响的研究却十分缺乏。以黄河三角洲芦苇(Phragmites australis)湿地生态系统为研究对象,将直径60 cm的圆形样方进行三个水平的石油添加处理(不添加石油或每年添加5 mm或10 mm厚的石油),同时通过切断或不切断样方内外芦苇根状茎的连接来控制克隆整合的有无。实验连续开展了两年(2014—2015年),每年8月份在样方内进行土壤样品取样,在实验室内采用磷酸脂肪酸(PLFA)法测定土壤微生物总量及主要微生物类群含量,并测定土壤微生物生物量碳和氮含量。取样时间显著影响土壤微生物PLFA总量、微生物碳和氮含量,这3个变量在2015年均显著高于2014年。石油添加在2015年显著增加了土壤微生物PLFA总量,但在2014年却无显著效应;同时,石油添加在2014年降低了土壤微生物碳和氮含量,而在2015年却增加了其含量。然而,无论在2014年还是2015年,芦苇的克隆整合对土壤微生物PLFA含量、微生物碳和氮含量均没有显著影响。土壤微生物PLFA总量与土壤微生物碳和氮含量呈正相关关系。这些结果表明,石油污染可以显著影响湿地土壤微生物动态,但克隆整合却无显著效应。     

An analysis of the costs and benefits of physiological integration between ramets in the clonal perennial herb Glechoma hederacea

Summary The costs and benefits, measured in terms of dry weight, of physiological integration between clonal ramets, were analysed in two experiments conducted on the clonal herb Glechoma hederacea. Firstly, integration between consecutively-produced ramets was examined in an experiment in which stolons grew from one set of growing conditions (either unshaded or shaded and either nutrient-rich or nutrient-poor) into conditions in which light or nutrient level was altered. Comparisons were made between the dry weight of the parts of the clones produced before and after growing conditions were changed, and the dry weights of the corresponding part of control clones subjected to constant growing conditions. In a second experiment, integration between two distinct parts of G. hederacea clones was investigated. In this experiment clones were grown from two connected parent ramets and the parts of the clone produced by each parent ramet were subjected independently to either nutrient-rich or nutrient-poor conditions. Ramets in resource-rich conditions provided considerable physiological support to those in resource-poor conditions. This was measured as a dry weight gain compared with the weight of the corresponding part of the control clones growing in resource-poor conditions. However, when stolons grew from resource-poor conditions into resource-rich conditions, there was no similar evidence of the resourcepoor ramtes receiving support from resource-rich ramets. Physiological integration did not result in dry weight gains when this would have necessitated basipetal translocation of resources.Because of the predominantly acropedal direction of movement of translocates in G. hederacea, the structure of the clone was important in determining the effectiveness of integration between ramets. Where physiological integration was effective, the cost to the supporting ramets in terms of dry weight was insignificant. Physiological integration allows clones to maintain a presence in less favourable sites with insignificant cost to ramets in favourable sites, thereby reducing the probability of invasion by other plants, and providing the potential for rapid clonal growth if conditions improve. Integrated support of ramets in unfavourable conditions also enables the clone to grow through unfavourable sites, thus increasing the probability of encountering more favourable conditions by wider foraging.     

Patch size and distance: modelling habitat structure from the perspective of clonal growth

Background and Aims

This study considers the spatial structure of patchy habitats from the perspective of plants that forage for resources by clonal growth. Modelling is used in order to compare two basic strategies, which differ in the response of the plant to a patch boundary. The ‘avoiding plant’ (A) never grows out of a good (resource-rich) patch into a bad (resource-poor) region, because the parent ramet withdraws its subsidy from the offspring. The ‘entering plant’ (E) always crosses the boundary, as the offspring is subsidized at the expense of the parent. In addition to these two extreme scenarios, an intermediate mixed strategy (M) will also be tested. The model is used to compare the efficiency of foraging in various habitats in which the proportion of resource-rich areas (p) is varied.

Methods

A stochastic cellular automata (CA) model is developed in which habitat space is represented by a honeycomb lattice. Each cell within the lattice can accommodate a single ramet, and colonization can occur from a parent ramet''s cell into six neighbouring cells. The CA consists of two layers: the population layer and the habitat. In the population layer, a cell can be empty or occupied by a ramet; in the habitat layer, a cell can be good (resource-rich) or bad (resource-poor). The habitat layer is constant; the population layer changes over time, according to the birth and death of ramets.

Key Results

Strategies M and E are primarily limited by patch distance, whereas A is more sensitive to patch size. At a critical threshold of the proportion of resource-rich areas, p = 0·5, the mean patch size increases abruptly. Below the threshold, E is more efficient than A, whilst above the threshold the opposite is true. The mixed strategy (M) is more efficient than either of the pure strategies across a broad range of p values.

Conclusions

The model predicts more species/genotypes with the ‘entering’ strategy, E, in habitats where resource-rich patches are scattered, and more plants with the ‘avoiding’ strategy, A, in habitats where the connectivity of resource-rich patches is high. The results suggest that the degree of physiological integration between a parent and an offspring ramet is important even across a very short distance because it can strongly influence the efficiency of foraging.     

Light limitation and litter of an invasive clonal plant,Wedelia trilobata,inhibit its seedling recruitment

Background and Aims

Invasive clonal plants have two reproduction patterns, namely sexual and vegetative propagation. However, seedling recruitment of invasive clonal plants can decline as the invasion process proceeds. For example, although the invasive clonal Wedelia trilobata (Asteraceae) produces numerous seeds, few seedlings emerge under its dense population canopy in the field. In this study it is hypothesized that light limitation and the presence of a thick layer of its own litter may be the primary factors causing the failure of seedling recruitment for this invasive weed in the field.

Methods

A field survey was conducted to determine the allocation of resources to sexual reproduction and seedling recruitment in W. trilobata. Seed germination was also determined in the field. Effects of light and W. trilobata leaf extracts on seed germination and seedling growth were tested in the laboratory.

Key Results

Wedelia trilobata blooms profusely and produces copious viable seeds in the field. However, seedlings of W. trilobata were not detected under mother ramets and few emerged seedlings were found in the bare ground near to populations. In laboratory experiments, low light significantly inhibited seed germination. Leaf extracts also decreased seed germination and inhibited seedling growth, and significant interactions were found between low light and leaf extracts on seed germination. However, seeds were found to germinate in an invaded field after removal of the W. trilobata plant canopy.

Conclusions

The results indicate that lack of light and the presence of its own litter might be two major factors responsible for the low numbers of W. trilobata seedlings found in the field. New populations will establish from seeds once the limiting factors are eliminated, and seeds can be the agents of long-distance dispersal; therefore, prevention of seed production remains an important component in controlling the spread of this invasive clonal plant.     

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.     

Short‐term fitness benefits of physiological integration in the clonal herb Hydrocotyle peduncularis

Abstract We test whether physiological integration enhances the short‐term fitness of the clonal herb Hydrocotyle peduncularis (Apiaceae, R. Brown ex A. Richards) subjected to spatial variation in water availability. Our measures of fitness and costs and benefits are based on the relative growth rate of fragmented genets. Physiological integration over a gradient in soil moisture resulted in a highly significant net benefit to genet growth of 0.015 g g^?1 day^?1. This net benefit represents a significant enhancement of the average fitness of fragmented genets spanning the moisture gradient relative to the average of those growing in homogeneous moist or dry conditions. Sections of genet fragments growing in dry conditions in spatially heterogeneous treatments had significantly higher growth than the sections they were connected to that were growing in moist conditions. Within fragments, older (parent) sections growing in moist conditions experienced significant costs from connection to younger (offspring) sections growing in dry conditions. In contrast, offspring sections with ample water did not experience any costs when connected to parent sections growing in dry conditions. However, the net benefit of physiological integration was similar for parent and offspring sections, suggesting that parent and offspring sections contributed equally to the net benefit of physiological integration to genet growth and short‐term fitness.     

Lack of genetic variation of an invasive clonal plant Eichhornia crassipes in China revealed by RAPD and ISSR markers

Random amplified polymorphic DNAs (RAPDs) and inter-simple sequence repeats (ISSRs) markers were used to analyze genetic structure of six populations of invasive plant Eichhornia crassipes that were sampled from its introduced regions in Southern China. Using 25 RAPD primers and 18 ISSR primers, 172 RAPD bands and 145 ISSR bands were produced respectively. But no polymorphic band was detected either within population or among populations by both markers, indicating the genetic diversity of E. crassipes in Southern China is extremely low, and all populations most likely consist of the same genotype. This study suggested that some other adaptability related factors, other than the genetic diversity, are responsible to the E. crassipes rapid expansion in China.     

Higher clonal integration in the facultative epiphytic fern Selliguea griffithiana growing in the forest canopy compared with the forest understorey

Background and Aims The advantage of clonal integration (resource sharing between connected ramets of clonal plants) varies and a higher degree of integration is expected in more stressful and/or more heterogeneous habitats. Clonal facultative epiphytes occur in both forest canopies (epiphytic habitats) and forest understories (terrestrial habitats). Because environmental conditions, especially water and nutrients, are more stressful and heterogeneous in the canopy than in the understorey, this study hypothesizes that clonal integration is more important for facultative epiphytes in epiphytic habitats than in terrestrial habitats.Methods In a field experiment, an examination was made of the effects of rhizome connection (connected vs. disconnected, i.e. with vs. without clonal integration) on survival and growth of single ramets, both young and old, of the facultative epiphytic rhizomatous fern Selliguea griffithiana (Polypodiaceae) in both epiphytic and terrestrial habitats. In another field experiment, the effects of rhizome connection on performance of ramets were tested in small (10 × 10 cm²) and large (20 × 20 cm²) plots in both epiphytic and terrestrial habitats.Key Results Rhizome disconnection significantly decreased survival and growth of S. griffithiana in both experiments. The effects of rhizome disconnection on survival of single ramets and on ramet number and growth in plots were greater in epiphytic habitats than in terrestrial habitats.Conclusions Clonal integration contributes greatly to performance of facultative epiphytic ferns, and the effects were more important in forest canopies than in forest understories. The results therefore support the hypothesis that natural selection favours genotypes with a higher degree of integration in more stressful and heterogeneous environments.     

Effects of fragmentation of clones compound over vegetative generations in the floating plant Pistia stratiotes

Background and AimsClonal plants dominate many plant communities, especially in aquatic systems, and clonality appears to promote invasiveness and to affect how diversity changes in response to disturbance and resource availability. Understanding how the special physiological and morphological properties of clonal growth lead to these ecological effects depends upon studying the long-term consequences of clonal growth properties across vegetative generations, but this has rarely been done. This study aimed to show how a key clonal property, physiological integration between connected ramets within clones, affects the response of clones to disturbance and resources in an aquatic, invasive, dominant species across multiple generations.MethodsSingle, parental ramets of the floating stoloniferous plant Pistia stratiotes were grown for 3 weeks, during which they produced two or three generations of offspring; connections between new ramets were cut or left intact. Individual offspring were then used as parents in a second 3-week iteration that crossed fragmentation with previous fragmentation in the first iteration. A third iteration yielded eight treatment combinations, zero to three rounds of fragmentation at different times in the past. The experiment was run once at a high and once at a low level of nutrients.ResultsIn each iteration, fragmentation increased biomass of the parental ramet, decreased biomass of the offspring and increased number of offspring. These effects persisted and compounded from one iteration to another, though more recent fragmentation had stronger effects, and were stronger at the low than at the high nutrient level. Fragmentation did not affect net accumulation of mass by groups after one iteration but increased it after two iterations at low nutrients, and after three iterations at both nutrient levels.ConclusionsBoth the positive and negative effects of fragmentation on clonal performance can compound and persist over time and can be stronger when resource levels are lower. Even when fragmentation has no short-term net effect on clonal performance, it can have a longer-term effect. In some cases, fragmentation may increase total accumulation of mass by a clone. The results provide the first demonstration of how physiological integration in clonal plants can affect fitness across generations and suggest that increased disturbance may promote invasion of introduced clonal species via effects on integration, perhaps especially at lower nutrient levels.     

Clonal integration supports the expansion from terrestrial to aquatic environments of the amphibious stoloniferous herb Alternanthera philoxeroides

Effects of clonal integration on land plants have been extensively studied, but little is known about the role in amphibious plants that expand from terrestrial to aquatic conditions. We simulated expansion from terrestrial to aquatic habitats in the amphibious stoloniferous alien invasive alligator weed ( Alternanthera philoxeroides ) by growing basal ramets of clonal fragments in soils connected (allowing integration) or disconnected (preventing integration) to the apical ramets of the same fragments submerged in water to a depth of 0, 5, 10 or 15 cm. Clonal integration significantly increased growth and clonal reproduction of the apical ramets, but decreased both of these characteristics in basal ramets. Consequently, integration did not affect the performance of whole clonal fragments. We propose that alligator weed possesses a double-edged mechanism during population expansion: apical ramets in aquatic habitats can increase growth through connected basal parts in terrestrial habitats; however, once stolon connections with apical ramets are lost by external disturbance, the basal ramets in terrestrial habitats increase stolon and ramet production for rapid spreading. This may contribute greatly to the invasiveness of alligator weed and also make it very adaptable to habitats with heavy disturbance and/or highly heterogeneous resource supply.     

Effects of soil nutrient heterogeneity on intraspecific competition in the invasive, clonal plant Alternanthera philoxeroides

Background and Aims

Fine-scale, spatial heterogeneity in soil nutrient availability can increase the growth of individual plants, the productivity of plant communities and interspecific competition. If this is due to the ability of plants to concentrate their roots where nutrient levels are high, then nutrient heterogeneity should have little effect on intraspecific competition, especially when there are no genotypic differences between individuals in root plasticity. We tested this hypothesis in a widespread, clonal species in which individual plants are known to respond to nutrient heterogeneity.

Methods

Plants derived from a single clone of Alternanthera philoxeroides were grown in the greenhouse at low or high density (four or 16 plants per 27·5 × 27·5-cm container) with homogeneous or heterogeneous availability of soil nutrients, keeping total nutrient availability per container constant. After 9 weeks, measurements of size, dry mass and morphology were taken.

Key Results

Plants grew more in the heterogeneous than in the homogeneous treatment, showing that heterogeneity promoted performance; they grew less in the high- than in the low-density treatment, showing that plants competed. There was no interactive effect of nutrient heterogeneity and plant density, supporting the hypothesis that heterogeneity does not affect intraspecific competition in the absence of genotypic differences in plasticity. Treatments did not affect morphological characteristics such as specific leaf area or root/shoot ratio.

Conclusions

Results indicate that fine-scale, spatial heterogeneity in the availability of soil nutrients does not increase competition when plants are genetically identical, consistent with the suggestion that effects of heterogeneity on competition depend upon differences in plasticity between individuals. Heterogeneity is only likely to increase the spread of monoclonal, invasive populations such as that of A. philoxeroides in China.     

Assessing the effects of land-use change on plant traits, communities and ecosystem functioning in grasslands: a standardized methodology and lessons from an application to 11 European sites

BACKGROUND AND AIMS: A standardized methodology to assess the impacts of land-use changes on vegetation and ecosystem functioning is presented. It assumes that species traits are central to these impacts, and is designed to be applicable in different historical, climatic contexts and local settings. Preliminary results are presented to show its applicability. METHODS: Eleven sites, representative of various types of land-use changes occurring in marginal agro-ecosystems across Europe and Israel, were selected. Climatic data were obtained at the site level; soil data, disturbance and nutrition indices were described at the plot level within sites. Sixteen traits describing plant stature, leaf characteristics and reproductive phase were recorded on the most abundant species of each treatment. These data were combined with species abundance to calculate trait values weighed by the abundance of species in the communities. The ecosystem properties selected were components of above-ground net primary productivity and decomposition of litter. KEY RESULTS: The wide variety of land-use systems that characterize marginal landscapes across Europe was reflected by the different disturbance indices, and were also reflected in soil and/or nutrient availability gradients. The trait toolkit allowed us to describe adequately the functional response of vegetation to land-use changes, but we suggest that some traits (vegetative plant height, stem dry matter content) should be omitted in studies involving mainly herbaceous species. Using the example of the relationship between leaf dry matter content and above-ground dead material, we demonstrate how the data collected may be used to analyse direct effects of climate and land use on ecosystem properties vs. indirect effects via changes in plant traits. CONCLUSIONS: This work shows the applicability of a set of protocols that can be widely applied to assess the impacts of global change drivers on species, communities and ecosystems.