Both phenotypic plasticity and locally adapted ecotypes may contribute to the success of invasive species in a wide range of habitats. Here, we conducted common garden experiments and molecular marker analysis to test the two alternative hypotheses in invasive alligator weed (Alternanthera philoxeroides), which colonizes both aquatic and terrestrial habitats. Ninety individuals from three pairs of aquatic versus terrestrial populations across southern China were analyzed, using inter simple sequence repeat (ISSR) marker, to examine population differentiation in neutral loci. Two common gardens simulating aquatic and terrestrial habitats were set up to examine population differentiation in quantitative traits. We found no evidence of population differentiation in both neutral loci and quantitative traits. Most individuals shared the same ISSR genotype. Meanwhile, plants from different habitats showed similar reaction norms across the two common gardens. In particular, plants allocated much more biomass to the belowground roots in the terrestrial environment, where alligator weed may lose part or all of the aboveground shoots because of periodical or accidental disturbances, than those in the aquatic environment. The combined evidence from molecular marker analysis and common garden experiments support the plasticity hypothesis rather than the ecotype hypothesis in explaining the adaptation of alligator weed in a wide range of habitats.
Phenotypic plasticity has been proposed as an important adaptive strategy for clonal plants in heterogeneous habitats. Increased phenotypic plasticity can be especially beneficial for invasive clonal plants, allowing them to colonize new environments even when genetic diversity is low. However, the relative importance of genetic diversity and phenotypic plasticity for invasion success remains largely unknown. Here, we performed molecular marker analyses and a common garden experiment to investigate the genetic diversity and phenotypic plasticity of the globally important weed Alternanthera philoxeroides in response to different water availability (terrestrial vs. aquatic habitats). This species relies predominantly on clonal propagation in introduced ranges. We therefore expected genetic diversity to be restricted in the two sampled introduced ranges (the USA and China) when compared to the native range (Argentina), but that phenotypic plasticity may allow the species' full niche range to nonetheless be exploited. We found clones from China had very low genetic diversity in terms of both marker diversity and quantitative variation when compared with those from the USA and Argentina, probably reflecting different introduction histories. In contrast, similar patterns of phenotypic plasticity were found for clones from all three regions. Furthermore, despite the different levels of genetic diversity, bioclimatic modeling suggested that the full potential bioclimatic distribution had been invaded in both China and USA. Phenotypic plasticity, not genetic diversity, was therefore critical in allowing A. philoxeroides to invade diverse habitats across broad geographic areas.
Summary1. Clonal plants benefit from the ability to translocate resources among interconnected ramets to colonize heterogeneous habitats. Clonal integration affects the resource level and morphological traits of ramets, and thus may influence their physiology and general performance. Although leaf gas exchange and its associated physiological adjustments are key traits to assess plant fitness, the effect of clonal integration on these functional traits is insufficiently understood. 2. In a glasshouse experiment, we addressed how clonal integration affects gas exchange properties, leaf characters and growth of ramets in two invasive plants, Alternanthera philoxeroides and Phyla canescens, under full sun and 85% shade. We also used stable-isotope labelling to assess the maternal subsidy to daughter ramets. 3. Similar effects of connection were observed in both species for most gas exchange parameters and leaf characters. Clonal integration did not affect photosynthetic capacity and respiratory rates of ramets. When grown in shade, ramets connected with an unshaded mother plant displayed higher area-based leaf nitrogen and chlorophyll content than severed ramets, but the additional nitrogen and chlorophyll was not translated to increased photosynthetic capability. Overall, severed ramets displayed significant light response for leaf nitrogen (area-based), photosynthetic nitrogen use efficiency, chlorophyll to nitrogen ratio, and nitrogen stable-isotope signature, but the light response was eliminated by clonal integration in connected ramets. 4. Both species displayed substantial maternal carbohydrate subsidy that benefited the growth of daughter ramets, but species-specific patterns were observed in the growth of daughter ramets and the amount of subsidy. The amount of subsidy was independent of ramet growth light levels for P. canescens, but shaded, connected ramets of A. philoxeroides received more subsidy than unshaded controls, facilitating a larger growth improvement relative to severed counterparts than P. canescens. 5. Synthesis. We observed increased leaf nitrogen and chlorophyll in shaded, connected ramets of two clonal invasive plants. Clonal integration may facilitate nitrogen assimilation and allow preacclimation to high-light conditions for shaded, connected ramets, thus promoting the opportunistic expansion of these invaders on site scale. The species-specific maternal subsidy pattern demonstrated that clonal plants possess different strategies to subsidize ramets under light-limited conditions.
Summary• Here, we compared the carbon isotope ratios of leaf respiratory CO 2 ( δ 13 C R ) and leaf organic components (soluble sugar, water soluble fraction, starch, protein and bulk organic matter) in five C 3 plants grown in a glasshouse and inside Biosphere 2. One species, Populus deltoides , was grown under three different CO 2 concentrations.• The Keeling plot approach was applied to the leaf scale to measure leaf δ 13 C R and these results were compared with the δ 13 C of leaf organic components.• In all cases, leaf respiratory CO 2 was more 13 C-enriched than leaf organic components. The amount of 13 C enrichment displayed a significant species-specific pattern, but the effect of CO 2 treatment was not significant on P. deltoides .• In C 3 plant leaves, 13 C-enriched respiratory CO 2 appears widespread. Among currently hypothesized mechanisms contributing to this phenomenon, non-statistical carbon isotope distribution within the sugar substrates seems most likely. However, caution should be taken when attempting to predict the δ 13 C of leaf respiratory CO 2 at the ecosystem scale by upscaling the relationship between leaf δ 13 C R and δ 13 C of leaf organic components.
Summary 1.Leaf respiratory temperature responses and general leaf properties of Quercus rubra were measured throughout the 2003 growing season in a deciduous forest in the northeastern USA. Measurements were made in the upper and lower portions of the canopy at two sites with different soil water availability. Correlations among respiration and various leaf properties were examined. 2. At a set temperature (10 and 20 ° C), area-based leaf respiration rates were higher in both the early and late growing season than in the mid-growing season (0·50 vs 0·33 µ mol CO 2 m − 2 s − 1 at 10 ° C, on average). Upper-canopy leaves generally had higher respiration rates than lower-canopy leaves (0·53 vs 0·30 µ mol CO 2 m − 2 s − 1 at 10 ° C, on average). At the drier site a more significant seasonal pattern in respiration was observed, while at the more mesic site a stronger canopy-position effect was detected. E 0 , a model variable related to the overall energy of activation of respiration, varied only slightly (52 ± 5 kJ mol − 1 K − 1 ), and was not influenced by season, site or canopy position. 3. Leaf properties (specific leaf area, nitrogen, soluble sugars) also varied with season, site and canopy position. Leaf N and reducing monose were positively correlated with leaf respiration rates. After isolating single factors (season, site, canopy position), reducing monose could partially explain the seasonality in respiration (32 − 79%), and leaf N ( N area ) was well correlated with the canopy-position effect. 4. Our results suggest that the temporal and spatial heterogeneities of respiration need to be considered in ecosystem models, but significant simplifications may be made in Q. rubra by assuming a constant temperature coefficient ( E 0 , 52·5 kJ mol − 1 in this study) or predicting the base respiration rate ( R 0 ) from well understood leaf properties.
Rapid adaptive evolution has been advocated as a mechanism that promotes invasion. Demonstrating adaptive evolution in invasive species requires rigorous analysis of phenotypic shifts driven by selection. Here, we document selection-driven evolution of Phyla canescens, an Argentine weed, in two invaded regions (Australia and France). Invasive populations possessed similar or higher diversity than native populations, and displayed mixed lineages from different sources, suggesting that genetic bottlenecks in both countries might have been alleviated by multiple introductions. Compared to native populations, Australian populations displayed more investment in sexual reproduction, whereas French populations possessed enhanced vegetative reproduction and growth. We partitioned evolutionary forces (selection vs. stochastic events) using two independent methods. Results of both analyses suggest that the pattern of molecular and phenotypic variability among regions was consistent with selection-driven evolution, rather than stochastic events. Our findings indicate that selection has shaped the evolution of P. canescens in two different invaded regions.
Early leafing and extended leaf longevity can be important mechanisms for the invasion of the forest understory. We compared the leaf phenology and photosynthetic characteristics of Berberis thunbergii, an early leafing invasive shrub, and two co-occurring native species, evergreen Kalmia latifolia and late leafing Vaccinium corymbosum, throughout the 2004 growing season. Berberis thunbergii leafed out 1 month earlier than V. corymbosum and approximately 2 weeks prior to the overstory trees. The photosynthetic capacity [characterized by the maximum carboxylation rate of Rubisco (V (cmax)) and the RuBP regeneration capacity mediated by the maximum electron transport rate (J (max))] of B. thunbergii was highest in the spring open canopy, and declined with canopy closure. The 2003 overwintering leaves of K. latifolia displayed high V (cmax) and J (max) in spring 2004. In new leaves of K. latifolia produced in 2004, the photosynthetic capacity gradually increased to a peak in mid-September, and reduced in late November. V. corymbosum, by contrast, maintained low V (cmax) and J (max) throughout the growing season. In B. thunbergii, light acclimation was mediated by adjustment in both leaf mass per unit area and leaf N on a mass basis, but this adjustment was weaker or absent in K. latifolia and V. corymbosum. These results indicated that B. thunbergii utilized high irradiance in the spring while K. latifolia took advantage of high irradiance in the fall and the following spring. By contrast, V. corymbosum generally did not experience a high irradiance environment and was adapted to the low irradiance understory. The apparent success of B. thunbergii therefore, appeared related to a high spring C subsidy and subsequent acclimation to varying irradiance through active N reallocation and leaf morphological modifications.
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