Alien balsams, strawberries and their pollinators in a warmer world.

BACKGROUND: Strawberries are a common crop whose yield success depends on the availability of pollinators. Invasive alien plants, such as Impatiens glandulifera and I. parviflora, are also attractive for bees and hoverflies, respectively, and occur in close proximity to strawberry cultivation areas. The aim of the study was to test whether alien plants may decrease pollination of strawberry cultivation. However, even if the pollinators are abundant, efficiency of their pollination may decrease as a result of revisits of flowers that were already probed. It is addressed by pollinators by scent marking. Moreover, such revisits can be determined by nectar replenishment, which may occur rapidly in nectar-rich flowers. We studied revisits to I. glandulifera by bumblebees and defined the factors that influence the probability of revisits (air temperature; pollinator species; family caste and size; flower area; sun radiation; and time of day). RESULTS: We found that the two alien species decreased the number of pollinators visiting strawberries. Apoidea, Bombini and Syrphidae significantly decreased on Fragaria × ananassa when alien Impatiens were present. We also revealed the influence of increasing air temperature on bumblebee foraging, which was particularly significant for female workers. At very high temperatures (> 37°C), bumblebee males revisited probed flowers less often than female workers. CONCLUSIONS: Our results demonstrate that in experimental conditions attractive alien species decrease pollination of strawberries, which may negatively affect production of this crop. Although the results have not been verified in real-life strawberry fields yet, we recommend that alien plant species that share the same pollinators and occur in close proximity of strawberries are controlled. Moreover, we found that revisits of probed flowers may weaken feeding efficiency of bumblebees. If revisits are not induced by nectar replenishment, then global warming may pose a serious threat to the survival of colonies, which may have consequences also for the plants that attract them, e.g., for strawberries.

The ability of seeds to float with water currents contributes to the invasion success of Impatiens balfourii and I. glandulifera.

Two alien species in Europe, Impatiens glandulifera and I. balfourii, are closely related, have similar growth rates and reproductive capacities, and are very attractive to pollinators. Nevertheless, only I. glandulifera is a highly invasive alien species in Europe, while I. balfourii is non-invasive. We assumed that the varying levels of invasiveness are driven by differences in the floating ability of their seeds, which may determine the invasion success of riparian alien plants, such as the Impatiens species. By mimicking two types of aquatic conditions, we determined seed floating ability for each species from younger and older populations. We also analyzed four seed traits: seed viability, surface, shape and coat structure. Seeds of the non-invasive I. balfourii float less well than seeds of the invasive I. glandulifera. We also found that the seeds of I. balfourii from the younger population have a higher floating ability in comparison with that of the seeds from the older population. The results for I. glandulifera were the opposite, with decreased floating ability in the younger population. These differences were associated with seed surface, shape and coat structure. These results indicate that the floating ability of I. balfourii seeds may increase over time following its introduction into a given area, while in the case of I. glandulifera, this ability may gradually decrease. Therefore, the former species, currently regarded as a poor disperser, has the potential to become invasive in the future, whereas the latter does not seem to benefit from further investments in the floating ability of its seeds.

Tolerance of Impatiens balsamina L., and Crotalaria retusa L. to grow on soil contaminated by used lubricating oil: A comparative study.

Screening of plant species with an ability to grow on contaminated soil is the most critical step in the planning of a phytoremediation program. While flourishing growth of Impatiens balsamina L. and Crotalaria retusa L. has been observed in areas adjacent to automobile service stations in Sri Lanka, no systematic study of their tolerance to used lubricating oil (ULO) contaminated soil has been carried out. Therefore, the aim of the present study was to investigate the comparative responses of I. balsamina L. and C. retusa L. to soil contaminated with ULO. Both species exhibited 100% seed germination in soils treated with 1%-5% w/w ULO. After 120 h exposure, root lengths and biomass of germinated seedlings of both species were significantly (p < 0.05) reduced in all treatments above 3% w/w ULO. The measured growth parameters of plants following 90 d exposure to 0.5-3% w/w ULO, indicated significant (p < 0.05) negative effects on I. balsamina and C. retusa at >1% w/w and >2% w/w ULO, respectively. There were no significant effects on chlorophyll content or root anatomy of either species under any treatments. Therefore, we concluded that I. balsamina can tolerate up to 1% of ULO and C. retusa up to 2% w/w ULO without displaying any negative effects. Comparatively higher biodegradation of ULO in the rhizosphere, root nodule formation, increases in root length and root hair density are all possible strategies for the exhibited higher tolerance of C. retusa. Therefore, the overall results indicate that C. retusa has the greater potential to be used in phytoremediation of ULO contaminated soils. The findings of the present study will be beneficial in planning phytoremediation program for ULO contaminated soil.

Drift happens: Molecular genetic diversity and differentiation among populations of jewelweed (Impatiens capensis Meerb.) reflect fragmentation of floodplain forests.

Landscape features often shape patterns of gene flow and genetic differentiation in plant species. Populations that are small and isolated enough also become subject to genetic drift. We examined patterns of gene flow and differentiation among 12 floodplain populations of the selfing annual jewelweed (Impatiens capensis Meerb.) nested within four river systems and two major watersheds in Wisconsin, USA. Floodplain forests and marshes provide a model system for assessing the effects of habitat fragmentation within agricultural/urban landscapes and for testing whether rivers act to genetically connect dispersed populations. We generated a panel of 12,856 single nucleotide polymorphisms and assessed genetic diversity, differentiation, gene flow, and drift. Clustering methods revealed strong population genetic structure with limited admixture and highly differentiated populations (mean multilocus FST  = 0.32, FST ' = 0.33). No signals of isolation by geographic distance or environment emerged, but alleles may flow along rivers given that genetic differentiation increased with river distance. Differentiation also increased in populations with fewer private alleles (R2  = 0.51) and higher local inbreeding (R2  = 0.22). Populations varied greatly in levels of local inbreeding (FIS  = 0.2-0.9) and FIS increased in more isolated populations. These results suggest that genetic drift dominates other forces in structuring these Impatiens populations. In rapidly changing environments, species must migrate or genetically adapt. Habitat fragmentation limits both processes, potentially compromising the ability of species to persist in fragmented landscapes.

Fertilizer effects of composted materials from different sources on cultivating Impatiens balsamina L. in municipal solid waste management.

At different stages of municipal solid waste management, several technologies such as home composting, industrial composting, and landfill mining could be used to recycle organic matters. Assessing the quality of composted material is crucial for determining where and how for recycling the organic fractions of municipal solid waste (OFMSW). Current studies mainly focused on comparing their biochemical characteristics and environmental impacts; however, comprehensive effects on cultivating plants were rarely compared with composted materials from different sources. Here, the final composting products from home composting (HC), industrial composting (IC), and landfill mining (LM), with different mixing ratios between OFMSW and soil (25, 50, 75, and 100%), were applied for cultivating Impatiens balsamina L. to examine the growing and flowering features under 195 days of observation. We found that all types of composted materials showed positive effects on growth of impatiens; however, their individual profiles were significant different. Generally, compost from HC showed the best comprehensive effects on the plant. Impatiens' dry weight biomass and maximum number of leaves and flowers of HC were1.5 and 2.8 times, 1.1 and 1.6 times, and 1.8 and 4.2 times than those of IC and LM, respectively. Compost from IC was superior in prolonging leaf-growing phase and increasing photosynthesis pigment contents of impatiens. Although comprehensive effect of fine fraction from landfill mining was much lower than HC and IC compost, it still improved impatiens growth and flowering compared to normal sandy soil. The results suggest that direct comprehensive effect on plants growth, flowering, and physiological influences could be introduced as an indicator when we compare different approach to recycle organics from MSW. Comprehensive effect on plants growth, flowering, and physiological influences could be introduced as a direct indicator for assessing organic waste recycling.

Biotic and abiotic drivers of intraspecific trait variation within plant populations of three herbaceous plant species along a latitudinal gradient.

BACKGROUND: The importance of intraspecific trait variation (ITV) is increasingly acknowledged among plant ecologists. However, our understanding of what drives ITV between individual plants (ITVBI) at the population level is still limited. Contrasting theoretical hypotheses state that ITVBI can be either suppressed (stress-reduced plasticity hypothesis) or enhanced (stress-induced variability hypothesis) under high abiotic stress. Similarly, other hypotheses predict either suppressed (niche packing hypothesis) or enhanced ITVBI (individual variation hypothesis) under high niche packing in species rich communities. In this study we assess the relative effects of both abiotic and biotic niche effects on ITVBI of four functional traits (leaf area, specific leaf area, plant height and seed mass), for three herbaceous plant species across a 2300 km long gradient in Europe. The study species were the slow colonizing Anemone nemorosa, a species with intermediate colonization rates, Milium effusum, and the fast colonizing, non-native Impatiens glandulifera. RESULTS: Climatic stress consistently increased ITVBI across species and traits. Soil nutrient stress, on the other hand, reduced ITVBI for A. nemorosa and I. glandulifera, but had a reversed effect for M. effusum. We furthermore observed a reversed effect of high niche packing on ITVBI for the fast colonizing non-native I. glandulifera (increased ITVBI), as compared to the slow colonizing native A. nemorosa and M. effusum (reduced ITVBI). Additionally, ITVBI in the fast colonizing species tended to be highest for the vegetative traits plant height and leaf area, but lowest for the measured generative trait seed mass. CONCLUSIONS: This study shows that stress can both reduce and increase ITVBI, seemingly supporting both the stress-reduced plasticity and stress-induced variability hypotheses. Similarly, niche packing effects on ITVBI supported both the niche packing hypothesis and the individual variation hypothesis. These results clearly illustrates the importance of simultaneously evaluating both abiotic and biotic factors on ITVBI. This study adds to the growing realization that within-population trait variation should not be ignored and can provide valuable ecological insights.

Disturbance response indicators of Impatiens walleriana exposed to benzene and chromium.

The purpose of this study was to evaluate the remediation potential and disturbance response indicators of Impatiens walleriana exposed to benzene and chromium. Numerous studies over the years have found abundant evidence of the carcinogenicity of benzene and chromium (VI) in humans. Benzene and chromium are two toxic industrial chemicals commonly found together at contaminated sites, and one of the most common management strategies employed in the recovery of sites contaminated by petroleum products and trace metals is in situ remediation. Given that increasing interest has focused on the use of plants as depollution agents, direct injection tests and benzene misting were performed on I. walleriana to evaluate the remediation potential of this species. I. walleriana accumulated hexavalent chromium, mainly in the root system (164.23 mg kg-1), to the detriment of the aerial part (39.72 mg kg-1), and presented visible damage only at the highest concentration (30 mg L-1). Unlike chromium (VI), chromium (III) was retained almost entirely by the soil, leaving it available for removal by phytotechnology. However, after the contamination stopped, I. walleriana responded positively to the detoxification process, recovering its stem stiffness and leaf color. I. walleriana showed visible changes such as leaf chlorosis during the ten days of benzene contamination. When benzene is absorbed by the roots, it is translocated to and accumulated in the plant's aerial part. This mechanism the plant uses ensures its tolerance to the organic compound, enabling the species to survive and reproduce after treatment with benzene. Although I. walleriana accumulates minor amounts of hexavalent chromium in the aerial part, this amount suffices to induce greater oxidative stress and to increase the amount of hydrogen peroxide when compared to that of benzene. It was therefore concluded that I. walleriana is a species that possesses desirable characteristics for phytotechnology.

Cadmium Uptake by Cuttings of Impatiens walleriana in Response to Different Cadmium Concentrations and Growth Periods.

Impatiens walleriana (I. walleriana), a potential cadmium (Cd) hyperaccumulator, can propagate by cuttings, which are less expensive to grow than seedlings. Different growth periods for cuttings, however, may lead to different physiological characteristics. In this study, I. walleriana cuttings were hydroponically grown in Cd-containing solutions (1.0-10.0 µM) for various growth periods (10-60 days). Experimental results showed that the Cd treatments had negative effects on growth compared to the controls that were not spiked with Cd. The extension of the growth period promoted most of the growth exhibitions of I. walleriana, except for SPAD readings for cuttings grown in the 5.0 and 10.0 µM solutions. The accumulation of Cd also increased over time, except in the roots of the cuttings grown in the 5.0 and 10.0 µM solutions. The subcellular distribution and chemical forms of Cd showed that I. walleriana developed better tolerance and detoxification capacities in the cuttings grown in the 5.0 and 10.0 µM solutions than in the cuttings grown in the other two Cd treatments.

Effects of extended growth periods on subcellular distribution, chemical forms, and the translocation of cadmium in Impatiens walleriana.

Impatiens walleriana plants accumulate sufficiently high concentrations of cadmium (Cd) for this species to be considered a potential Cd hyperaccumulator. Rooted cuttings were grown hydroponically for 25 and 50 days in solutions spiked with various Cd concentrations. The subcellular distribution and chemical forms of Cd in different organs were analyzed, and its upward translocation was also assessed. The plants accumulated large amounts of Cd; the Cd concentration in the roots and shoots reached 120-1900 and 60-1600 mg/kg, respectively. Regardless of the growth period, the Cd accumulated in the roots was primarily compartmentalized in the soluble fraction or ethanol and deionized water extractable chemical forms with high migration abilities. Translocation to the shoots was followed by an association of Cd mainly in the cell wall or with pectate and protein. The roots' Cd showed a high migration capacity for predicting the shoots' Cd concentrations. Different exposure periods significantly affected the subcellular distribution of Cd in the stems, and thus the upward translocation.

An invasive plant alters phenotypic selection on the vegetative growth of a native congener.

PREMISE OF THE STUDY: The ecological consequences of plant competition have frequently been tested, but the evolutionary outcomes of these interactions have gone largely unexplored. The study of species invasions can make an important contribution to this field of research by allowing us to watch ecological and evolutionary processes unfold as a novel species is integrated into a plant community. We explored the ecological and evolutionary impact of an invasive jewelweed, Impatiens glandulifera, on a closely related native congener, I. capensis and asked: (1) Does the presence of the invasive jewelweed alter the fitness of native jewelweed populations? (2) Does the invasive jewelweed affect the vegetative growth of the native congener? and (3) Does the invasive jewelweed alter phenotypic selection on the vegetative traits of the native congener? METHODS: We used a greenhouse competition experiment, an invasive species removal field experiment, and a survey of natural populations. KEY RESULTS: We show that when the invasive jewelweed is present, phenotypic selection favors native jewelweed individuals investing less in rapid upward growth and more in branching and fruiting potential through the production of nodes. CONCLUSIONS: This research demonstrates that invasive plants have the potential to greatly alter natural selection on native competitors. Studies investigating altered selection in invaded communities can reveal the potential evolutionary impact of invasive competitors, while deepening our understanding of the more general role of competition in driving plant evolution and permitting species coexistence.

Utilization of solar energy in sewage sludge composting: fertilizer effect and application.

Three reactors, ordinary, greenhouse, and solar, were constructed and tested to compare their impacts on the composting of municipal sewage sludge. Greenhouse and solar reactors were designed to evaluate the use of solar energy in sludge composting, including their effects on temperature and compost quality. After 40 days of composting, it was found that the solar reactor could provide more stable heat for the composting process. The average temperature of the solar reactor was higher than that of the other two systems, and only the solar reactor could maintain the temperature above 55°C for more than 3 days. Composting with the solar reactor resulted in 31.3% decrease in the total organic carbon, increased the germination index to 91%, decreased the total nitrogen loss, and produced a good effect on pot experiments.

More cells, bigger cells or simply reorganization? Alternative mechanisms leading to changed internode architecture under contrasting stress regimes.

Shading and mechanical stress (MS) modulate plant architecture by inducing different developmental pathways. Shading results in increased stem elongation, often reducing whole-plant mechanical stability, while MS inhibits elongation, with a concomitant increase in stability. Here, we examined how these organ-level responses are related to patterns and processes at the cellular level by exposing Impatiens capensis to shading and MS. Shading led to the production of narrower cells along the vertical axis. By contrast, MS led to the production of fewer, smaller and broader cells. These responses to treatments were largely in line with genetic differences found among plants from open and closed canopy sites. Shading- and MS-induced plastic responses in cellular characteristics were negatively correlated: genotypes that were more responsive to shading were less responsive to MS and vice versa. This negative correlation, however, did not scale to mechanical and architectural traits. Our data show how environmental conditions elicit distinctly different associations between characteristics at the cellular level, plant morphology and biomechanics. The evolution of optimal response to different environmental cues may be limited by negative correlations of stress-induced responses at the cellular level.

Effect of intra- and interspecific competition on the performance of native and invasive species of Impatiens under varying levels of shade and moisture.

Many alien plants are thought to be invasive because of unique traits and greater phenotypic plasticity relative to resident species. However, many studies of invasive species are unable to quantify the importance of particular traits and phenotypic plasticity in conferring invasive behavior because traits used in comparative studies are often measured in a single environment and by using plants from a single population. To obtain a deeper insight into the role of environmental factors, local differences and competition in plant invasions, we compared species of Impatiens (Balsaminaceae) of different origin and invasion status that occur in central Europe: native I. noli-tangere and three alien species (highly invasive I. glandulifera, less invasive I. parviflora and potentially invasive I. capensis). In two experiments we harvested late-stage reproductive plants to estimate performance. The first experiment quantified how populations differed in performance under varying light and moisture levels in the absence of competition. The second experiment quantified performance across these environments in the presence of intra- and inter-specific competition. The highly invasive I. glandulifera was the strongest competitor, was the tallest and produced the greatest biomass. Small size and high plasticity were characteristic for I. parviflora. This species appeared to be the second strongest competitor, especially under low soil moisture. The performance of I. capensis was within the range of the other Impatiens species studied, but sometimes limited by alien competitors. Our results suggest that invasion success within the genus Impatiens depends on the ability to grow large under a range of environmental conditions, including competition. The invasive species also exhibited greater phenotypic plasticity across environmental conditions than the native species. Finally, the decreased performance of the native I. noli-tangere in competition with other species studied indicates that this species may be possibly excluded from its sites by invading congeners.

Effect of methionine on production of naphthoquinones in Impatiens balsamina root cultures and detection of some secondary metabolites.

CONTEXT: Lawsone, lawsone methyl ether and 3,3'-methylelnebislawsone are the main active compounds of Impatiens balsamina L. (Balsaminaceae). These compounds possess various pharmacological activities that have been shown to assist with the treatment of skin diseases. OBJECTIVE: This work focused on increased naphthoquinone production in I. basamina root cultures using methionine feeding. MATERIALS AND METHODS: I. balsamina root cultures were maintained in liquid Gamborg's B5 medium supplemented with 0.1 mg/L α-naphthalene acetic acid, 0.1 mg/L kinetin, 1.0 mg/L 6-benzyladenine and 20 g/L sucrose. The effect of methionine concentration (50, 100, 300, 500 and 1000 mg/L) on naphthoquinone production of I. basamina root cultures was determined. Isolation of secondary metabolites from I. balsamina root cultures was also carried out. RESULTS AND DISCUSSION: Feeding of 300 mg/L methionine to the root cultures at the beginning of the growth cycle increased the production of 3,3'-methylelnebislawsone almost two-fold (0.63 mg/g dry weight, compared to the control group 0.32 mg/g dry weight). Optimization of the feeding conditions showed that adding 500 mg/L methionine to a 21-day old root cultures increased production of lawsone methyl ether and 3,3'-methylenebislawsone up to 2.6- and 3.1-fold higher, respectively, compared to the controls. In addition, various pharmacologically interesting secondary metabolites were isolated from I. balsamina root cultures, such as a flavonoid, luteolin, a naphthoquinone, 2,3-dihydroxy-1,4-naphthoquinone, and a triterpenoid, echinocystic acid. This is the first report of the occurrence of these compounds in this plant.

What is the impact of Impatiens parviflora on diversity and composition of herbal layer communities of temperate forests?

The aim was to estimate the impacts of invasive Impatiens parviflora on forests' herbal layer communities. A replicated Before-After-Control-Impact field experiment and comparisons with adjacent uninvaded plots were used. The alien's impact on species richness was tested using hierarchical generalized mixed effect models with Poisson error structure. Impact on species composition was tested using multivariate models (DCA, CCA, RDA) and Monte-Carlo permutation tests. Removal plots did not differ in native species richness from neither invaded nor adjacent uninvaded plots, both when the treatment's main effect or its interaction with sampling time was tested (Chi(2) = 0.4757, DF = 2, p = 0.7883; Chi(2) = 7.229, DF = 8, p = 0.5121 respectively). On the contrary, ordination models revealed differences in the development of plots following the treatments (p = 0.034) with the invaded plots differing from the adjacent uninvaded (p = 0.002). Impatiens parviflora is highly unlikely to impact native species richness of invaded communities, which may be associated with its limited ability to create a dense canopy, a modest root system or the fact the I. parviflora does not represent a novel and distinctive dominant to the invaded communities. Concerning its potential impacts on species composition, the presence of native clonal species (Athyrium filix-femina, Dryopteris filix-mas, Fragaria moschata, Luzula luzuloides, Poa nemoralis) on the adjacent uninvaded plots likely makes them different from the invaded plots. However, these competitive and strong species are more likely to prevent the invasion of I. parviflora on the adjacent uninvaded plots rather than being themselves eliminated from the invaded communities.

Effects of CO(2) enrichment on growth and development of Impatiens hawkeri.

The effects of CO(2) enrichment on growth and development of Impatiens hawkeri, an important greenhouse flower, were investigated for the purpose of providing scientific basis for CO(2) enrichment to this species in greenhouse. The plants were grown in CO(2)-controlled growth chambers with 380 (the control) and 760 (CO(2) enrichment) µmol · mol(-1), respectively. The changes in morphology, physiology, biochemistry, and leaf ultrastructure of Impatiens were examined. Results showed that CO(2) enrichment increased flower number and relative leaf area compared with the control. In addition, CO(2) enrichment significantly enhanced photosynthetic rate, contents of soluble sugars and starch, activities of peroxidase (POD), superoxide dismutase (SOD), and ascorbate peroxidase (APX), but reduced chlorophyll content and malondialdehyde (MDA) content. Furthermore, significant changes in chloroplast ultrastructure were observed at CO(2) enrichment: an increased number of starch grains with an expanded size, and an increased ratio of stroma thylakoid to grana thylakoid. These results suggest that CO(2) enrichment had positive effects on Impatiens, that is, it can improve the visual value, promote growth and development, and enhance antioxidant capacity.

Bacterial reduction of Cr(VI) at technical scale--the Malaysian experience.

The bacterial reduction of Cr(VI) from industrial wastewater was evaluated using a 2.0-m(3) bioreactor. Liquid pineapple waste was used as a nutrient for the biofilm community formed inside the bioreactor. The use of rubber wood sawdust as packing material was able to immobilize more than 10(6) CFU mL(-1) of Acinetobacter haemolyticus cells after 3 days of contact time. Complete reduction of 15-240 mg L(-1) of Cr(VI) was achieved even after 3 months of bioreactor operation. Cr(VI) was not detected in the final effluent fraction indicating complete removal of Cr from solution from the flocculation/coagulation step and the unlikely re-oxidation of Cr(III) into Cr(VI). Impatiens balsamina L. and Gomphrena globosa L. showed better growth in the presence of soil-sludge mixture compared to Coleus scutellarioides (L.) Benth. Significant amounts of Cr accumulated at different sections of the plants indicate its potential application in Cr phytoremediation effort. The bacterial-based system was also determined not to be detrimental to human health based on the low levels of Cr detected in the hair and nail samples of the plant operators. Thus, it can be said that bacterial-based Cr(VI) treatment system is a feasible alternative to the conventional system especially for lower Cr(VI) concentrations, where sludge generated can be used as growth supplement for ornamental plant as well as not detrimental to the health of the workers.

Seedling traits, plasticity and local differentiation as strategies of invasive species of Impatiens in central Europe.

BACKGROUND AND AIMS: Invasiveness of some alien plants is associated with their traits, plastic responses to environmental conditions and interpopulation differentiation. To obtain insights into the role of these processes in contributing to variation in performance, we compared congeneric species of Impatiens (Balsaminaceae) with different origin and invasion status that occur in central Europe. METHODS: Native I. noli-tangere and three alien species (highly invasive I. glandulifera, less invasive I. parviflora and potentially invasive I. capensis) were studied and their responses to simulated canopy shading and different nutrient and moisture levels were determined in terms of survival and seedling traits. KEY RESULTS AND CONCLUSIONS: Impatiens glandulifera produced high biomass in all the treatments and the control, exhibiting the 'Jack-and-master' strategy that makes it a strong competitor from germination onwards. The results suggest that plasticity and differentiation occurred in all the species tested and that along the continuum from plasticity to differentiation, the species at the plasticity end is the better invader. The most invasive species I. glandulifera appears to be highly plastic, whereas the other two less invasive species, I. parviflora and I. capensis, exhibited lower plasticity but rather strong population differentiation. The invasive Impatiens species were taller and exhibited higher plasticity and differentiation than native I. noli-tangere. This suggests that even within one genus, the relative importance of the phenomena contributing to invasiveness appears to be species' specific.

The multiple fuzzy origins of woodiness within Balsaminaceae using an integrated approach. Where do we draw the line?

BACKGROUND AND AIMS: The family Balsaminaceae is essentially herbaceous, except for some woodier species that can be described as 'woody' herbs or small shrubs. The family is nested within the so-called balsaminoid clade of Ericales, including the exclusively woody families Tetrameristaceae and Marcgraviaceae, which is sister to the remaining families of the predominantly woody order. A molecular phylogeny of Balsaminaceae is compared with wood anatomical observations to find out whether the woodier species are derived from herbaceous taxa (i.e. secondary woodiness), or whether woodiness in the family represents the ancestral state for the order (i.e. primary woodiness). METHODS: Wood anatomical observations of 68 Impatiens species and Hydrocera triflora, of which 47 are included in a multigene phylogeny, are carried out using light and scanning electron microscopy and compared with the molecular phylogenetic insights. KEY RESULTS: There is much continuous variation in wood development between the Impatiens species studied, making the distinction between herbaceousness and woodiness difficult. However, the most woody species, unambiguously considered as truly woody shrubs, all display paedomorphic wood features pointing to secondary woodiness. This hypothesis is further supported by the molecular phylogeny, demonstrating that these most woody species are derived from herbaceous (or less woody) species in at least five independent clades. Wood formation in H. triflora is mostly confined to the ribs of the stems and shows paedomorphic wood features as well, suggesting that the common ancestor of Balsaminaceae was probably herbaceous. CONCLUSIONS: The terms 'herbaceousness' and 'woodiness' are notoriously difficult to use in Balsaminaceae. However, anatomical observations and molecular sequence data show that the woodier species are derived from less woody or clearly herbaceous species, demonstrating that secondary woodiness has evolved in parallel.

Testing mechanisms and context dependence of costs of plastic shade avoidance responses in Impatiens capensis (Balsaminaceae).

PREMISE OF THE STUDY: Many plant species elongate their shoots in response to neighbor proximity and neighbor height. Although these plastic responses may be beneficial in terms of enhancing light interception, they also may have costs in terms of increased risk of mechanical failure (i.e., lodging or breaking) because of thinner stems. This trade-off between light acquisition and stability may shape the evolution of plastic elongation responses to foliage shade. METHODS: In a field experiment manipulating elongation phenotypes and densities, we tested two hypotheses. We predicted that the risks of mechanical failure depend on plastic elongation and/or on characteristics of the immediate neighborhood, such as density and neighbor height. Further, we predicted that plants that fail mechanically would have reduced fitness. KEY RESULTS: Mechanical failure was earlier and more frequent at high density and showed a complex interaction with neighborhood characteristics such as relative height of the neighbors and the expression of early plasticity. Plants that broke earlier had shorter lifespan and lower reproductive output. CONCLUSIONS: Our results show that depending on the height and density of the group, plastic elongation responses can remain advantageous despite costs of increased risk of mechanical failure of the taller stems, as mechanical failure was not associated with strong costs in terms of reduced lifespan or seed production. The overall benefits of elongation outweigh the costs resulting in selection for elongation at the population level.