Fei-Hai Yu

United we stand, divided we fall: a meta-analysis of experiments on clonal integration and its relationship to invasiveness.

Many ecosystems are dominated by clonal plants. Among the most distinctive characteristics of clonal plants is their potential for clonal integration (i.e. the translocation of resources between interconnected ramets), suggesting that integration may play a role in their success. However, a general synthesis of effects of clonal integration on plant performance is lacking. We conducted a meta-analysis on the effects of clonal integration on biomass production and asexual reproduction of the whole clone, the recipient part (i.e. the part of a clone that imports resources) and the donor part (i.e. the part of a clone that exports resources). The final dataset contained 389 effect sizes from 84 studies covering 57 taxa. Overall, clonal integration increased performance of recipient parts without decreasing that of donor parts, and thus increased performance of whole clones. Among the studies and taxa considered, the benefits of clonal integration did not differ between two types of experimental approaches, between stoloniferous and rhizomatous growth forms, between directions of resource translocation (from younger to older ramet or vice versa), or among types of translocated resources (water, nutrients and carbohydrates). Clonal taxa with larger benefits of integration on whole-clone performance were not more invasive globally, but taxa in which recipient parts in unfavorable patches benefited more from integration were. Our results demonstrate general performance benefits of clonal integration, at least in the short term, and suggest that clonal integration contributes to the success of clonal plants.

Clonal Integration Affects Growth, Photosynthetic Efficiency and Biomass Allocation, but not the Competitive Ability, of the Alien Invasive Alternanthera philoxeroides under Severe Stress

BACKGROUND AND AIMS Many notorious alien invasive plants are clonal, but little is known about some roles and aspects of clonal integration. Here, the hypothesis is tested that clonal integration affects growth, photosynthetic efficiency, biomass allocation and competitive ability of the exotic invasive weed Alternanthera philoxeroides (Amaranthaceae). METHODS The apical parts of Alternanthera were grown either with or without the lawn grass Schedonorus phoenix (tall fescue) and their stolon connections to the basal parts grown without competitors were either severed or left intact. KEY RESULTS Competition greatly reduced the maximum quantum yield of photosystem II (F(v)/F(m)) and growth (biomass, number of ramets and leaves, total stolon length and total leaf area) of the apical Alternanthera, but not the biomass of S. phoenix. Stolon connections significantly increased F(v)/F(m) and growth of Alternanthera. However, such effects on growth were smaller with than without competition and stolon connections did not alter the relative neighbour effect of Alternanthera. Stolon connections increased Alternantheras biomass allocation to roots without competition, but decreased it with competition. CONCLUSIONS Clonal integration contributed little to Alternantheras competitive ability, but was very important for Alternanthera to explore open space. The results suggest that the invasiveness of Alternanthera may be closely related to clonal integration.

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.

Adaptation of Rhizome Connections in Drylands: Increasing Tolerance of Clones to Wind Erosion

BACKGROUND AND AIMS Wind erosion is a severe stress for plants in drylands, but the mechanisms by which plants withstand erosion remain largely unknown. Here, the hypothesis is tested that maintaining rhizome connections helps plants to tolerate erosion. METHODS Five transects were established across an inland dune in Inner Mongolia, China, and measurements were made of leaf number, biomass per ramet and rhizome depth of Psammochloa villosa in 45 plots. In 40 x 40 cm plots of P. villosa on another dune, the top 15 or 30 cm of sand was removed for 1.5 or 3 months to simulate short- and long-term moderate and severe erosion, respectively, with untreated plots as controls, and the rhizomes at the edges of half of the plots were severed to mimic loss of rhizome connections. KEY RESULTS Leaf number and biomass per ramet showed quadric relationships with rhizome depth; when rhizomes were exposed to the air, the associated ramets either died or became very weak. Ramet number, leaf number and biomass per plot decreased with increasing erosion severity. Rhizome connections did not affect these traits under control or short-term erosion, but increased them under long-term erosion. CONCLUSIONS Rhizome connections alleviated the negative effects of erosion on P. villosa, very likely because the erosion-stressed ramets received water and/or photosynthates translocated from those connected ramets that were not subject to erosion. This study provides the first evidence that maintaining rhizome connections helps plants to tolerate erosion in drylands.

Physiological integration in an introduced, invasive plant increases its spread into experimental communities and modifies their structure

What determines the invasiveness of introduced plants is still poorly known. Many of the most invasive plant species are clonal, and physiological integration between connected individuals (ramets) of clonal plants may contribute to their ability to spread into communities and reduce performance of existing species. This contribution of integration to the invasiveness of clonal plants may be greater in denser communities. A greenhouse study was conducted to test these two hypotheses. High- and low-density communities were created by sowing seeds of eight grassland species. Each community was planted with three ramets of the stoloniferous, introduced plant Alternanthera philoxeroides that were disconnected from or left connected to ramets growing on bare soil. Connection increased the spread of Alternanthera within a community, but did not reduce community biomass. Alternanthera grew less in high-density communities, but connection did not improve its growth more than in low-density communities. Low-density communities had higher evenness when Alternanthera was connected than when it was disconnected because shoot mass was lower in the more abundant species in the community and higher in the less abundant ones. These results partly supported the first hypothesis, but not the second. The effect of integration on community structure could be due to higher resource import by the ramets of Alternanthera closer to the dominant species. Integration therefore can increase the initial spread of new clonal plant species into communities and modify the effects of this spread on community structure.

Effects of fragmentation on the survival and growth of the invasive, clonal plant Alternanthera philoxeroides

The capacity of small clonal fragments to survive and grow is a major factor in the spread of clonal plants, especially aquatic species. This study of the introduced, invasive, amphibious, stoloniferous herb Alternanthera philoxeroides in China tested the effects of fragment size, of loss of the stolon apex, and of the original position of a fragment within a clone before fragmentation on the survival and growth of fragments in a greenhouse experiment. The stolon internodes of plants consisting of one stolon with 5 ramets were severed in all 16 possible patterns, generating fragments of 1–5 ramets. These 16 fragmentation treatments were crossed with removal of the stolon apex. Fragmentation and apex removal did not affect total growth of plants, but more highly fragmented plants produced smaller, more numerous ramets. The survival of fragments consisting of one original ramet was about 85%, compared to 100% survival of fragments with 5 original ramets. Fragments consisting of the youngest ramets along the original stolon grew more than fragments of the same size that consisted of older ramets. These effects of fragmentation on growth provide new evidence for clonal integration of the rate at which connected ramets produce new ramets and novel evidence that fragments produced from different portions of clones have different potential for spread. They further suggest that detachment of small groups of distal ramets along stolons in the field might be very effective in spreading the species and that a trade-off between the number and the size of new ramets produced by fragments may represent an adaptive, plastic response to disturbance. Results do not recommend intentional fragmentation of clones as a means of controlling the rate of growth of this invasive species but suggest that incidental fragmentation may not increase its growth in mass, although it may increase its rate of spread.

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.

Effects of denudation and burial on growth and reproduction of Artemisia ordosica in Mu Us sandland

Plants in a dune environment have a high risk of being denudated by wind or buried by sand. We conducted a field experiment to assess growth and reproductive performance as well as biomass allocation of Artemisia ordosica after denudation and burial. Height growth decreased after denudation, but remained constant after burial; biomass was reduced by severe burial, but not by moderate ones. Vegetative growth measured by current-year generated vegetative branches declined with increasing severity of burial. Reproductive maturity was not affected by either denudation or burial, and reproduction decreased only after severe denudation. After denudation, biomass allocated to reproduction was greatly reduced, as shown by strongly reduced biomass allocation to fruits and whole reproductive branches, but that allocated to current-year vegetative growth was maintained. Biomass allocation to reproduction and vegetative growth remained rather stable after burial. Current-year reproductive-to-vegetative-biomass ratio was lower after denudation than after burial, indicating that vegetative growth was more favored by A. ordosica after denudation. These modifications are adaptive in the fluctuating environment of mobile sand dunes where denudation and burial frequently occur.

Trampling, defoliation and physiological integration affect growth, morphological and mechanical properties of a root-suckering clonal tree

BACKGROUND AND AIMS Grazing is a complex process involving the simultaneous occurrence of both trampling and defoliation. Clonal plants are a common feature of heavily grazed ecosystems where large herbivores inflict the simultaneous pressures of trampling and defoliation on the vegetation. We test the hypothesis that physiological integration (resource sharing between interconnected ramets) may help plants to deal with the interactive effects of trampling and defoliation. METHODS In a field study, small and large ramets of the root-suckering clonal tree Populus simonii were subjected to two levels of trampling and defoliation, while connected or disconnected to other ramets. Plant responses were quantified via survival, growth, morphological and stem mechanical traits. KEY RESULTS Disconnection and trampling increased mortality, especially in small ramets. Trampling increased stem length, basal diameter, fibrous root mass, stem stiffness and resistance to deflection in connected ramets, but decreased them in disconnected ones. Trampling decreased vertical height more in disconnected than in connected ramets, and reduced stem mass in disconnected ramets but not in connected ramets. Defoliation reduced basal diameter, leaf mass, stem mass and leaf area ratio, but did not interact with trampling or disconnection. CONCLUSIONS Although clonal integration did not influence defoliation response, it did alleviate the effects of trampling. We suggest that by facilitating resource transport between ramets, clonal integration compensates for trampling-induced damage to fine roots.

Invasive clonal plant species have a greater root-foraging plasticity than non-invasive ones.

Clonality is frequently positively correlated with plant invasiveness, but which aspects of clonality make some clonal species more invasive than others is not known. Due to their spreading growth form, clonal plants are likely to experience spatial heterogeneity in nutrient availability. Plasticity in allocation of biomass to clonal growth organs and roots may allow these plants to forage for high-nutrient patches. We investigated whether this foraging response is stronger in species that have become invasive than in species that have not. We used six confamilial pairs of native European clonal plant species differing in invasion success in the USA. We grew all species in large pots under homogeneous or heterogeneous nutrient conditions in a greenhouse, and compared their nutrient-foraging response and performance. Neither invasive nor non-invasive species showed significant foraging responses to heterogeneity in clonal growth organ biomass or in aboveground biomass of clonal offspring. Invasive species had, however, a greater positive foraging response in terms of root and belowground biomass than non-invasive species. Invasive species also produced more total biomass. Our results suggest that the ability for strong root foraging is among the characteristics promoting invasiveness in clonal plants.