Phenotypic plasticity is commonly considered as a trait associated with invasiveness in alien plants because it may enhance the ability of plants to occupy a wide range of environments. Although the evidence of greater phenotypic plasticity in invasive plants is considerable, it is not yet conclusive. We used a meta‐analysis approach to evaluate whether invasive plant species show greater phenotypic plasticity than their native or non‐invasive counterparts. The outcome of such interspecific comparisons may be biased when phylogenetic relatedness is not taken into account. Consequently, species pairs belonged to the same genus, tribe or family. The meta‐analysis included 93 records from 35 studies reporting plastic responses to light, nutrients, water, CO2, herbivory and support availability. Contrary to what is often assumed, overall, phenotypic plasticity was similar between invasive plants and native or non‐invasive closely related species. The same result was found when separate analyses were conducted for trait plasticity to nutrients, light and water availability. Thus, invasive plant species and their native or non‐invasive counterparts are equally capable of displaying functional responses to environmental heterogeneity. The colonization of a wide range of environments by invasive plants could be due to their capacity to undergo adaptive ecotypic differentiation rather than to their ability to display plastic responses. Alternatively, phenotypic plasticity might play a role in plant invasion, but only during the initial phases, when tolerance of the novel environment is essential for plant survival. Afterwards, once alien plants are identified as invaders, the magnitude of phenotypic plasticity might be reduced after selection of the optimum phenotypes in each habitat. The identification of plant traits that consistently predict invasiveness might be a futile task because different traits favor invasiveness in different environments. Approaches at the local scale, focusing on the ecology of specific invasive plants, could be more fruitful than global macro‐analyses.
Adaptation to heterogeneous environments can occur via phenotypic plasticity, but how often this occurs is unknown. Reciprocal transplant studies provide a rich dataset to address this issue in plant populations because they allow for a determination of the prevalence of plastic versus canalized responses. From 31 reciprocal transplant studies, we quantified the frequency of five possible evolutionary patterns: (1) canalized response–no differentiation: no plasticity, the mean phenotypes of the populations are not different; (2) canalized response–population differentiation: no plasticity, the mean phenotypes of the populations are different; (3) perfect adaptive plasticity: plastic responses with similar reaction norms between populations; (4) adaptive plasticity: plastic responses with parallel, but not congruent reaction norms between populations; and (5) nonadaptive plasticity: plastic responses with differences in the slope of the reaction norms. The analysis included 362 records: 50.8% life-history traits, 43.6% morphological traits, and 5.5% physiological traits. Across all traits, 52% of the trait records were not plastic, and either showed no difference in means across sites (17%) or differed among sites (83%). Among the 48% of trait records that showed some sort of plasticity, 49.4% showed perfect adaptive plasticity, 19.5% adaptive plasticity, and 31% nonadaptive plasticity. These results suggest that canalized responses are more common than adaptive plasticity as an evolutionary response to environmental heterogeneity.
Phenotypic plasticity is essential for plant adaptation to changing environments but some factors limit its expression, causing plants to fail in producing the best phenotype for a given environment. Phenotypic integration refers to the pattern and magnitude of character correlations and it might play a role as an internal constraint to phenotypic plasticity. We tested the hypothesis that phenotypic integration Á estimated as the number of significant phenotypic correlations between traits Á constrains phenotypic plasticity of plants. The rationale is that, for any phenotypic trait, the more linked with other traits it is, the more limited is its range of variation. In the perennial species Convolvulus chilensis (Convolvulaceae) and Lippia alba (Verbenaceae) we determined the relationship between phenotypic plasticity to relevant environmental factors Á shading for C. chilensis and drought for L. alba Á and the magnitude of phenotypic integration of morphological and biomass allocation traits. In C. chilensis plants, plasticity to shading of a given trait decreased with the number of significant correlations that it had with the other traits. Likewise, the characters that showed greater plasticity to experimental drought in L. alba plants had fewer significant phenotypic correlations with other characters. We report a novel limit to phenotypic plasticity of plants by showing that the phenotypic trait architecture may constrain their plastic, functional responses to the environment.
Charophytes represent the group of green algae whose ancestors invaded land and ultimately gave rise to land plants 450 million years ago. While Zygnematophyceae are believed to be the direct sister lineage to embryophytes, different members of this group ( Penium , Spirogyra , Zygnema ) and the advanced thallus forming Coleochaete as well as the sarcinoid basal streptophyte Chlorokybus were investigated concerning their vegetative extracellular matrix (ECM) properties. Many taxa exhibit adhesion phenomena that are critical for affixing to a substrate or keeping cells together in a thallus, however, there is a great variety in possible reactions to e.g., wounding. In this study an analysis of adhesion mechanisms revealed that arabinogalactan proteins (AGPs) are most likely key adhesion molecules. Through use of monoclonal antibodies (JIM13) or the Yariv reagent, AGPs were located in cell surface sheaths and cell walls that were parts of the adhesion focal zones on substrates including wound induced rhizoid formation. JIM5, detecting highly methyl-esterfied homoglacturonan and JIM8, an antibody detecting AGP glycan and LM6 detecting arabinans were also tested and a colocalization was found in several examples (e.g., Zygnema ) suggesting an interplay between these components. AGPs have been described in this study to perform both, cell to cell adhesion in algae forming thalli and cell to surface adhesion in the filamentous forms. These findings enable a broader evolutionary understanding of the function of AGPs in charophyte green algae.
The extracellular matrix (ECM) of many charophytes, the assemblage of green algae that are the sister group to land plants, is complex, produced in large amounts, and has multiple essential functions. An extensive secretory apparatus and endomembrane system are presumably needed to synthesize and secrete the ECM, but structural details of such a system have not been fully characterized. Penium margaritaceum is a valuable unicellular model charophyte for studying secretion dynamics. We report that Penium has a highly organized endomembrane system, consisting of 150–200 non-mobile Golgi bodies that process and package ECM components into different sets of vesicles that traffic to the cortical cytoplasm, where they are transported around the cell by cytoplasmic streaming. At either fixed or transient areas, specific cytoplasmic vesicles fuse with the plasma membrane and secrete their constituents. Extracellular polysaccharide (EPS) production was observed to occur in one location of the Golgi body and sometimes in unique Golgi hybrids. Treatment of cells with brefeldin A caused disruption of the Golgi body, and inhibition of EPS secretion and cell wall expansion. The structure of the endomembrane system in Penium provides mechanistic insights into how extant charophytes generate large quantities of ECM, which in their ancestors facilitated the colonization of land.
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