Clonal Transgenerational Effects Transmit for Multiple Generations in a Floating Plant

Environmental conditions of a parent plant can influence the performance of their clonal offspring, and such clonal transgenerational effects may help offspring adapt to different environments. However, it is still unclear how many vegetative generations clonal transgenerational effects can transmit for and whether it depends on the environmental conditions of the offspring. We grew the ancestor ramets of the floating clonal plant Spirodela polyrhiza under a high and a low nutrient level and obtained the so-called 1^st-generation offspring ramets of two types (from these two environments). Then we grew the 1^st-generation offspring ramets of each type under the high and the low nutrient level and obtained the so-called 2^nd-generation offspring ramets of four types. We repeated this procedure for another five times and analyzed clonal transgenerational effects on growth, morphology and biomass allocation of the 1^st- to the 6^th-generation offspring ramets. We found positive, negative or neutral (no) transgenerational effects of the ancestor nutrient condition on the offspring of S. polyrhiza, depending on the number of vegetative generations, the nutrient condition of the offspring environment and the traits considered. We observed significant clonal transgenerational effects on the 6^th-generation offspring; such effects occurred for all three types of traits (growth, morphology and allocation), but varied depending on the nutrient condition of the offspring environment and the traits considered. Our results suggest that clonal transgenerational effects can transmit for multiple vegetative generations and such impacts can vary depending on the environmental conditions of offspring.     

Seed Banks and Species Richness Potential of Coal Slurry Ponds Reclaimed as Wetlands

Seed bank experiments are described to assess the species richness potential of coal slurry ponds reclaimed as wetlands (ranging from 6 to more than 40 years old). Experimental treatments test the drawdown and flooded conditions characteristic of the vegetation dynamics of emergent wetlands in the Upper Mississippi Valley. More seedlings, primarily annuals, emerged from exposed wet sediments (freely drained) than under continuous flooded sediments in cold ponds (339 versus 136 seedlings m^?2, respectively) and in natural ponds (163 versus 47, respectively). More seeds were produced by plants established in freely drained conditions than under flooded conditions from sediments in the coal ponds (26546 versus 1842 seeds m^?2, respectively) and the natural ponds (28430 versus 4526, respectively). Similarly, more biomass was also produced by these plants in freely drained than under flooded conditions in coal ponds (118 versus 47 g m^?2, respectively) and natural ponds (118 versus 52, respectively). Fertilization (NPK) did not affect germination for the most part, but it did affect seed set and biomass production, especially for C₄ annuals such as Echinochloa crusgalli and Panicum dichotomiflorum. I propose that lime (calcium carbonate) and fertilizer be applied during the first few scheduled drawdowns for these coal slurry ponds reclaimed as wetlands to increase the number of species and to allow their more rapid development as self-sustaining systems.     

Responses of the saltmarsh rush Juncus kraussii to salinity and waterlogging

Juncus kraussii Hochst., an important saltmarsh macrophyte, is intensively harvested for many commercially orientated products and current populations are under threat of overexploitation. In saline, intertidal mud banks, this species occurs on higher ground, suggesting that it is adapted to lower salinities and less frequent inundation. The objectives of this study were to determine biomass accumulation, as well as morphological and physiological adaptations of J. kraussii to salinity and waterlogging stresses. Plants collected from the field were subjected to 0.2, 10, 30, 50 and 70% seawater under drained or flooded conditions for three months. Measurements were made of biomass accumulation, CO₂ exchange, chlorophyll fluorescence, ion and water relations. Furthermore, seed germination responses to a range of salinities were investigated. Total dry biomass accumulation, as well as the number and height of culms, decreased with increase in salinity under both flooded and drained conditions. Generally, CO₂ exchange, stomatal conductance, Photosystem II (PSII) quantum yield and electron transport rate (ETR) through PSII declined with increase in salinity in both the flooded and drained treatments. Predawn and midday ψ in culms decreased with increase in salinity, being lower under drained than flooded conditions. Inorganic solute concentrations in culms increased with increase in salinity, with Na⁺ and Cl⁻ being the predominant ions. Na⁺/K⁺ ratios in culms increased significantly with increase in salinity. Proline concentrations in roots and culms, which increased with salinity, were considerably higher under drained than flooded conditions. Germination was best at salinities less than 20% seawater and decreased significantly with further increase in salinity to 110% seawater. Transfer of ungerminated salt-treated seeds to distilled water stimulated germination. This study has demonstrated that J. kraussii is a highly salt and flood tolerant species, being able to grow and survive in salinities up to 70% seawater, under both drained and flooded conditions. Maximal growth occurred at low salinities (<10% seawater) under flooded condition.     

Relationship between morphological and physiological responses to waterlogging and salinity in Sporobolus virginicus (L.) Kunth

The effects of waterlogging and salinity on morphological and physiological responses in the marsh grass Sporobolus virginicus (L.) Kunth were investigated in a 4×2 factorial experiment. Plants were subjected to four salinity levels (0, 100, 200 and 400 mol m^–3 NaCl) and two soil inundation conditions (drained and flooded) for 42 days. Flooding at 0 mol m^–3 NaCl caused initiation of adventitious surface roots, increased internal acration and plant height, induced alcohol dehydrogenase activity (ADH), and decreased belowground biomass and the number of culms per plant. Salinity increase from 0 to 400 mol m^–3 NaCl under drained conditions increased leaf and root proline concentrations and decreased photosynthesis, aboveground biomass, number of culms per plant and number of internodes per culm. Concurrent waterlogging and salinity induced ADH activity and adventitious surface roots but decreased plant height and aboveground biomass. Internal air space increased with waterlogging from 0 to 100 mol m^–3 NaCl but further increases in salinity to 400 mol m^–3 reduced air space. Combined waterlogging and salinity stresses, however, had no effect on photosynthesis or on the concentrations of proline in leaves or roots. These results are discussed in relation to the widespread colonization by S. virginicus of a wide range of coastal environments varying in soil salinity and in the frequency and intensity of waterlogging.     

Waterlogging responses of Schoenoplectus scirpoides (Schrad) Browning (Cyperaceae)

Waterlogging responses of an emergent freshwater macrophyte, Schoenoplectus scirpoides, were investigated by carrying out laboratory and field experiments. Soil cores containing uniform plants of S. scirpoides were subjected to drained, 1/3 flooded, alternately flooded and drained at weekly intervals, and continuously flooded treatments for 6 months. Mean soil redox potential of the drained soil was 225 ± 29 mV and was significantly lower in the other treatments. Total dry biomass, above‐ground biomass and number of culms in the flooded/drained and flooded treatments were significantly higher than those in the drained and 1/3 flooded treatments. Below‐ground/above‐ground biomass ratios were highest in the drained treatment and significantly lower in the waterlogged treatments. Plant height and the growth rate of culms were least in the drained treatment and significantly higher in the other treatments. Culm specific gravity decreased significantly with increase in soil moisture. In the field study, soil redox potentials were moderately reduced in the inland and streamside sites, while culm elongation rate was significantly higher by 94% in the streamside compared with the inland site. Optimum growth occurred under flooded conditions, suggesting that the species could flourish with increasing water intrusion in estuarine areas, as predicted by global climate change.     

To share or not to share: clonal integration in a submerged macrophyte in response to light stress

The ability of clonal plant species to share resources has been studied in many experiments. The submerged macrophyte Potamogeton perfoliatus produces interconnected ramets within short time intervals and hence may or may not share resources with ramets growing in less favourable microhabitats. From a genet point of view, sharing with ramets growing under less favourable conditions might not be an optimal strategy when photosynthates could be used to establish other ramets growing under more favourable conditions. To analyse the plasticity in clonal integration of P. perfoliatus, we set up a factorial aquaria experiment with unshaded or shaded recipient ramets (offspring), which were connected to or separated from donor ramets (parents). Increased biomass production of offspring in parent–offspring systems compared with severed offspring in both light and shade showed that ramets share resources through clonal integration. The relative translocation to the first- and second-offspring generation was influenced by habitat quality: If first-offspring ramets grew in a shaded microhabitat, second-offspring ramets clearly profited. This may be at least partially because of the fact that resources are shifted from first-offspring to second-offspring ramets, indicating controlled senescence of the first-offspring. This complex sharing behaviour might be relevant when plants produce ramets within a dense patch of macrophytes, where support of a shaded ramet might not pay off.     

Exploitation of patchy soil water resources by the clonal vine <Emphasis Type="Italic">Ficus tikoua</Emphasis> in karst habitats of southwestern China

The karst habitats of southwestern China are characterized by a highly heterogeneous distribution of water resources. We hypothesized that the clonal integration between connected ramets of the clonal vine Ficus tikoua was an important adaptive strategy to the patchy distribution of water resources in these habitats. We grew ramet pairs (each consisting of a parent and an offspring ramet) in both homogeneously and heterogeneously watered conditions. The offspring ramets were well-watered, whereas their connected parent ramets were randomly assigned to four water treatments: well-watered, mild water stress, moderate water stress, and severe water stress. Increasing water stress decreased leaf water potential, relative water content, net assimilation rate, maximum quantum yield of PSII (F _v/F _m), and biomass of the parent ramets. Subjecting the parents to water stress significantly increased root biomass and root mass ratio (RMR) of their offspring ramets. Exploitation of plentiful water resources through the increased adventitious roots connected to another soil patch permitted the complete restoration of water relations and photosynthetic capacity of offspring ramets after an initial depression. Water relations and gas exchange of the parents were not affected by the water supply to their connected offspring ramets, suggesting that offspring ramets hardly exported water to the stressed parents. However, net assimilation rate and proline content of the offspring ramets increased when they were connected to water-stressed parents. The compensatory photosynthetic responses of offspring ramets connected to stressed parents revealed an increasing trend as the experiment progressed. Morphological and physiological plasticity of F. tikoua in response to heterogeneous water resources allow them to adapt to karst habitats and be suitable candidates for vegetation restoration projects.     

Effects of clonal fragmentation on intraspecific competition of a stoloniferous floating plant

Disturbance is common and can fragment clones of plants. Clonal fragmentation may affect the density and growth of ramets so that it could alter intraspecific competition. To test this hypothesis, we grew one (low density), five (medium density) or nine (high density) parent ramets of the floating invasive plant Pistia stratiotes in buckets, and newly produced offspring ramets were either severed (with fragmentation) or remained connected to parent ramets (no fragmentation). Increasing density reduced biomass of the whole clone (i.e. parent ramet plus its offspring ramets), showing intense intraspecific competition. Fragmentation decreased biomass of offspring ramets, but increased biomass of parent ramets and the whole clone, suggesting significant resource translocation from parent to offspring ramets when clones were not fragmented. There was no interaction effect of density x fragmentation on biomass of the whole clone, and fragmentation did not affect competition intensity index. We conclude that clonal fragmentation does not alter intraspecific competition between clones of P. stratiotes, but increases biomass production of the whole clone. Thus, fragmentation may contribute to its interspecific competitive ability and invasiveness, and intentional fragmentation should not be recommended as a measure to stop the rapid growth of this invasive species.     

Flooding effects on rapid responses of the invasive plant Alternanthera philoxeroides to defoliation

In experiments under controlled growth conditions it was examined how flooding affected the responses of the invasive plant Alternanthera philoxeroides to defoliation. In drained and flooded conditions, plants were subjected to five defoliation levels: 0, 10, 50, 90% removal of leaf tissue and apex removal (90% leaf tissue plus apical bud removal). Plants were harvested weekly for five weeks. In drained conditions, plant biomasses including total biomass, shoot biomass and root biomass after 50% defoliation rapidly recovered to the control plant level. They were significantly lower for the 90% defoliation and apex removal treatments compared to control plants throughout the experiment. In flooded conditions, total biomass and shoot biomass after 50% defoliation, 90% defoliation, and apex removal treatments could return to control plant levels before the end of the experiment. In 90% defoliation and apex removal treatments root to shoot biomass ratios of both drained and flooded plants were initially much higher than in control plants, but the difference disappeared rapidly. The final biomasses decreased with increased defoliation intensity in drained conditions, but no significant difference was generally found in any of the defoliation treatments in flooded conditions. The rapid re-growth of A. philoxeroides plants after defoliation may partly be responsible for its invasion success. However, defoliation capable of removing 90% of the leaf tissue may be desirable in restricting the growth of this invasive species in drained conditions.     

Evolution of an invasive phenotype: shift to belowground dominance and enhanced competitive ability in the introduced range

In response to novel selection pressures in an introduced range, non-native species may evolve more competitive phenotypes unique from those of their native range. We examined the existence of an invasive phenotype in the herbaceous perennial Artemisia vulgaris, a frequent invader of the Northeast and Mid-Atlantic US. Populations from both the native (European) and the introduced (North American) ranges were grown in intra-specific competition (same population), inter-specific competition with the native perennial herb Solidago canadensis, and alone in a common garden to quantify shifts in resource allocation and neighbor effects on performance and competitive ability. Without competition, introduced A. vulgaris populations were much shorter than native populations, but germinated earlier, produced more ramets, more belowground and total biomass, and maintained higher root-to-shoot ratios. Under inter- and intra-specific competitions, introduced A. vulgaris populations were shorter, but produced more ramets, belowground, and total biomass than native populations. S. canadensis belowground and total biomass were more highly suppressed by introduced than native A. vulgaris. Our data suggest that since the introduction to North America, A. vulgaris has evolved a more competitive invasive phenotype characterized by many short ramets with more extensive root/rhizome networks. This rapid evolutionary shift likely benefits A. vulgaris in its introduced range by allowing establishment and subsequent dominance in dense stands of existing vegetation.     

Effects of Water Table Drawdown on Root Production and Aboveground Biomass in a Boreal Bog

We studied the effect of long-term water table drawdown on the vascular plant community in an ombrotrophic bog in central Finland by measuring aboveground biomass and belowground production (by in-growth cores) across plant functional groups including herbs, shrubs, and trees. We compared drained and undrained portions 45 years after the installation of a drainage ditch network, which has lowered water levels of 15–20 cm on average in the drained part of the site. Although shrub fine root production did not differ significantly between sites, water table drawdown increased belowground tree fine root production by 740% (3.8 ± 5.4 SD and 28.1 ± 24.1 g m^?2 y^?1 in undrained and drained sites, respectively) at the expense of herb root production, which declined 38% (27.62 ± 16.40 and 10.58 ± 15.7 g m^?2 y^?1 in undrained and drained sites, respectively) yielding no significant overall change in total fine root production. Drainage effects on aboveground biomass showed a similar pattern among plant types, as aboveground tree biomass increased dramatically with drainage (79 ± 135 and 2546 ± 1551 g m^?2 in drained and undrained sites, respectively). Although total shrub biomass was not significantly different between sites, shrubs allocated more biomass to stems than leaves in the drained site. Drainage also caused a significant shift in shrub species composition. Although trees dominated the aboveground biomass following water table drawdown, understorey vegetation, mainly shrubs, continued to dominate belowground fine root production, comprising 64% of total root production at the drained site. Aboveground biomass proved to be a good predictor of belowground production, suggesting that allometric relationships can be developed to estimate belowground production in these systems. Increase in tree root production can counteract decrease in herb fine root production following water table drawdown, emphasizing the importance of plant functional type responses to water table drawdown. Whether these changes will offset ecosystem C loss via increased plant C storage or stimulate soil organic matter decomposition via increased above- and belowground litter inputs requires further study.     

Genetic mosaics of ecosystem functioning across aspen-dominated landscapes

Genetic diversity is the foundation of all biodiversity, and the genetic variation within species is increasingly recognized as being important to ecosystem level processes. Recent research demonstrates that plant genotype influences above- and belowground communities as well as basic ecosystem functions. However, the extent to which plant genotypes create spatial mosaics of genetically mediated ecosystem processes in natural forests is uncertain. We use Populus tremuloides as a model system to demonstrate the importance of plant genotype on carbon and nitrogen cycling in natural systems. We identified 24 distinct P. tremuloides clones with multiple ramets across 25 km² in southern Wisconsin, United States, using microsatellite makers. We then sampled clone leaf chemistry and belowground nutrient content and microbial extracellular enzyme activity. Aspen-induced variation in belowground carbon and nitrogen content, and microbial activity, varied widely among clones. Variation in green leaf chemistry and belowground microbial activity were correlated with genetic distance among clones, such that more genetically distant clones created more divergent patches of ecosystem processes. These data suggest that aspen genotypes create spatial mosaics of genetically mediated ecosystem functioning across natural landscapes and can therefore have evolutionary consequences for co-occurring species.     

The effects of physiological integration on biomass partitioning in plant modules: an experimental study with the stoloniferous herb Glechoma hederacea

Morphological and physiological plasticity are crucial attributes enabling plants to acquire resources from heterogeneous habitats. Although physiological integration can modify biomass partitioning in modules, especially when connected modules experience different conditions, its ecological importance has been largely overlooked. This experiment examined its effects on above- and belowground biomass partitioning by modules in the stoloniferous herb Glechoma hederacea. We studied how biomass allocation to roots by younger ramets was affected by connection to older ramets, and by nutrient conditions. A lower proportion of biomass was allocated to roots by younger ramets growing under low nutrient (LN) conditions when connected to older ramets in high nutrient (HN) conditions than when they were isolated, demonstrating localised modification of biomass partitioning due to physiological integration. The proportion of biomass allocated to roots by younger ramets was also lower when connected to older ramets in HN conditions than when connected to older ramets in LN conditions. Thus, the effect of integration on biomass partitioning depended on the nutrient conditions experienced by connected ramets. Such changes in biomass partitioning would result in more extensive stolon growth, and greater lateral displacement of new ramets. Understanding the ecological implications of phenotypic plasticity in plants will require further examination of the effects of physiological integration when connected modules experience contrasting growing conditions. This study demonstrates that such integration affects the biomass allocation strategy of connected ramets, enhancing resource acquisition in heterogeneous habitats. The widespread success of clonality in many communities is likely to be strongly promoted by this characteristic.     

Belowground interactions shift the relative importance of direct and indirect genetic effects

Intraspecific genetic variation can affect decomposition, nutrient cycling, and interactions between plants and their associated belowground communities. However, the effects of genetic variation on ecosystems can also be indirect, meaning that genes in a focal plant may affect ecosystems by altering the phenotype of interacting (i.e., neighboring) individuals. We manipulated genotype identity, species identity, and the possibility of belowground interactions between neighboring Solidago plants. We hypothesized that, because our plants were nitrogen (N) limited, the most important interactions between focal and neighbor plants would occur belowground. More specifically, we hypothesized that the genotypic identity of a plant's neighbor would have a larger effect on belowground biomass than on aboveground biomass, but only when neighboring plants were allowed to interact belowground. We detected species‐ and genotype‐level variation for aboveground biomass and ramet production. We also found that belowground biomass and ramet production depended on the interaction of neighbor genotype identity and the presence or absence of belowground interactions. Additionally, we found that interspecific indirect genetic effects (IIGEs; changes in focal plant traits due to the genotype identity of a heterospecific neighbor) had a greater effect size on belowground biomass than did focal genotype; however, this effect only held in pots that allowed belowground interactions. These results expand the types of natural processes that can be attributed to genotypes by showing that, under certain conditions, a plant's phenotype can be strongly determined by the expression of genes in its neighbor. By showing that IIGEs are dependent upon plants being able to interact belowground, our results also provide a first step for thinking about how genotype‐based, belowground interactions influence the evolutionary outcomes of plant‐neighbor interactions.     

Nutrient addition does not increase the benefits of clonal integration in an invasive plant spreading from open patches into plant communities

• One benefit of clonal integration is that resource translocation between connected ramets enhances the growth of the ramets grown under stressful conditions, but whether such resource translocation reduces the performance of the ramets grown under favourable conditions has not produced consistent results. In this study, we tested the hypothesis that resource translocation to recipient ramets may reduce the performance of donor ramets when resources are limiting but not when resources are abundant.
• We grew Mikania micrantha stolon fragments (each consisting of two ramets, either connected or not connected) under spatially heterogeneous competition conditions such that the developmentally younger, distal ramets were grown in competition with a plant community and the developmentally older, proximal ramets were grown without competition. For half of the stolon fragments, slow‐release fertiliser pellets were applied to both the distal and proximal ramets.
• Under both the low and increased soil nutrient conditions, the biomass, leaf number and stolon length of the distal ramets were higher, and those of the proximal ramets were lower when the stolon internode was intact than when it was severed. For the whole clone, the biomass, leaf number and stolon length did not differ between the two connection treatments. Connection did not change the biomass of the plant communities competing with distal ramets of M. micrantha.
• Although clonal integration may promote the invasion of M. micrantha into plant communities, resource translocation to recipient ramets of M. micrantha will induce a cost to the donor ramets, even when resources are relatively abundant.

     

TRANSPORT OF CARBON AMONG CONNECTED RAMETS OF EICHHORNIA CRASSIPES (PONTEDERIACEAE) AT NORMAL AND HIGH LEVELS OF CO2

The floating, stoloniferous plant, Eichhornia crassipes, has high rates of productivity and rapidly invades new sites. Because the transport of carbon among connected ramets is known to increase the growth of clonal plants, we asked whether there is intraclonal carbon transport in E. crassipes. Because net photosynthesis of E. crassipes is significantly higher at high levels of atmospheric CO₂, we also asked if high CO₂ can change patterns of carbon transport in ways that might modify clonal growth. We exposed individual ramets within groups of connected ramets to ¹⁴CO₂ for 15–45 min and measured the distribution of ¹⁴C in the group after 4 days of growth at 350, 700, 1,400, or 2,800 μ1 1^−-1 CO₂. At 350 μ1 1^−-1 CO₂, a parent ramet exported approximately 10% of the ¹⁴C that it assimilated to its first rooted offspring ramet. The offspring exported a similar percentage of the ^l4C it assimilated toward the parent; two-thirds of this ¹⁴C was retained by the parent, and one-third moved into new offspring of the parent. In all ramets, imported carbon moved into leaves as well as roots. At the higher levels of CO₂, the percentage of assimilated carbon exported from a parent ramet to the leaf blades of its first offspring was lower by half. High CO₂ had little other effect on carbon transport. E. crassipes maintains bidirectional transport of carbon between ramets even under uniform and favorable environmental conditions and when external CO₂ levels are very high.     

Effects of size and growth rate on vegetative reproduction in Typha

Summary The objective of this study was to separate the effects of plant biomass and growth rate on vegetative reproduction in two species of cat-tail, Typha latifolia and T. angustifolia. Replicate clones of both species were grown under conditions of 100%, 42%, 24%, and 9% full sunlight with harvests at 41, 70, and 91 days after shading. T. angustifolia produced most of its vegetative offspring before the first harvest and increased biomass over the remainder of the experiment by increasing the size of its ramets. In contrast, T. latifolia produced vegetative offspring gradually throughout the experiment adding new ramets only after existing clones were of mature size. As a result of these differences in the cloning process, T. angustifolia showed little correlation between vegetative reproduction and clone size while T. latifolia showed a strong correlation between gegetative reproduction and clone size at the three highest light intensities. Growth rates, average clone size and vegetative reproduction were all reduced by reductions in light intensity for both species. However, no effect of growth rate on the relationship between clone size and vegetative reproduction in T. latifolia could be detected. T. latifolia showed greater survivorship and more biomass production under 9% light than T. angustifolia indicating a greater shade tolerance.     

Miscanthus × giganteus and Arundo donax shoot and rhizome tolerance of extreme moisture stress

Crops grown for bioenergy production are a mandated component of the United States energy portfolio. Giant miscanthus (Miscanthus × giganteus) is a leading bioenergy crop similar in habit to the invasive plant giant reed (Arundo donax). To characterize the environmental tolerance of giant miscanthus, we compared the soil moisture stress tolerance of giant miscanthus and giant reed under glasshouse conditions. We subjected both species to soil moisture conditions of severe drought (?4.2 MPa), mild drought (?0.5 MPa), field‐capacity (control), and flooded soils. These conditions were applied to two cohorts: one in which soil moisture conditions were imposed on newly planted rhizome fragments, and one in which conditions were imposed on established plants after 8 weeks of growth in field‐capacity soil. After 16 weeks, we harvested all plants, measured above‐ and belowground biomass, and evaluated the reproductive viability of rhizome fragments. The total biomass of each species under flooded conditions was not different from the field‐capacity control groups regardless of cohort. However, drought did affect the two cohorts differently. In the cohort treated after 8 weeks of growth, mild and severe drought conditions resulted in 56% and 66% reductions in biomass, averaged over both species, compared with the controls. In the cohort treated for the entire 16 weeks, mild and severe drought conditions resulted in 92% and 94% reductions in biomass. Rhizome fragments from both species and both cohorts showed 100% viability following flooded and control treatments; drought treatments reduced rhizome viability in both species, with a greater impact on giant miscanthus. Although giant miscanthus does not appear to have the potential to escape and establish in relatively dry upland ecosystems, it does show tolerance to flooded conditions similar to giant reed.     

Habitat selection in spatially heterogeneous environments: a test of foraging behaviour in the clonal submerged macrophyte Vallisneria spiralis

1. To test whether clonal macrophytes can select favourable habitats in heterogeneous environments, clonal fragments of the stoloniferous submerged macrophyte Vallisneria spiralis were subjected to conditions in which light intensity and substratum nutrients were patchily distributed. The allocation of biomass accumulation and ramet production of clones to the different patches was examined. 2. The proportion of both biomass and ramet number of clones allocated to rich patches was significantly higher than in poor patches. The greatest values of both clone and leaf biomass were produced in the heterogeneous light treatment, in which clones originally grew from light‐rich to light‐poor patches, while clones produced the most offspring ramets in the treatments with heterogeneous substratum nutrients. Similarly, root biomass had the highest values in nutrient‐rich patches when clones grew from nutrient‐rich to nutrient‐poor patches. 3. The quality of patches in which parent ramets established significantly influenced the foraging pattern. When previously established in rich patches, a higher proportion of biomass was allocated to rich patches, whereas a higher proportion of ramet number was allocated to rich patches when previously established in poor patches. 4. Results demonstrate that the clonal macrophyte V. spiralis can exhibit foraging in submerged heterogeneous environments: when established under resource‐rich conditions V. spiralis remained in favourable patches, whereas if established in adverse conditions it could escape by allocating more ramets to favourable patches.     

Effects of parental light environment on growth and morphological responses of clonal offspring

• Environments experienced by parent ramets of clonal plants can potentially influence fitness of clonal offspring ramets. Such clonal parental effects may result from heritable epigenetic changes, such as DNA methylation, which can be removed by application of DNA de‐methylation agents such as 5‐azacytidine.
• To test whether parental shading effects occur via clonal generation and whether DNA methylation plays a role in such effects, parent plants of the clonal herb Alternanthera philoxeroides were first subjected to two levels of light intensity (high versus low) crossed with two levels of DNA de‐methylation (no or with de‐methylation by application of 5‐azacytidine), and then clonal offspring taken from each of these four types of parent plant were subjected to the same two light levels.
• Parental shading effects transmitted via clonal generation decreased growth and modified morphology of clonal offspring. Offspring responses were also influenced by DNA methylation level of parent plants. For clonal offspring growing under low light, parental shading effects on growth and morphology were always negative, irrespective of the parental de‐methylation treatment. For clonal offspring growing under high light, parental shading effects on offspring growth and morphology were negative when the parents were not treated with 5‐azacytidine, but neutral when they were treated with 5‐azacytidine.
• Overall, parental shading effects on clonal offspring performance of A. philoxeroides were found, and DNA methylation is likely to be involved in such effects. However, parental shading effects contributed little to the tolerance of clonal offspring to shading.