The effects of osmotic stress on the cell wall-plasma membrane domains of the unicellular streptophyte, Penium margaritaceum.

Penium margaritaceum is a unicellular zygnematophyte (basal Streptophyteor Charophyte) that has been used as a model organism for the study of cell walls of Streptophytes and for elucidating organismal adaptations that were key in the evolution of land plants.. When Penium is incubated in sorbitol-enhance medium, i.e., hyperosmotic medium, 1000-1500 Hechtian strands form within minutes and connect the plasma membrane to the cell wall. As cells acclimate to this osmotic stress over time, further significant changes occur at the cell wall and plasma membrane domains. The homogalacturonan lattice of the outer cell wall layer is significantly reduced and is accompanied by the formation of a highly elongate, "filamentous" phenotype. Distinct peripheral thickenings appear between the CW and plasma membrane and contain membranous components and a branched granular matrix. Monoclonal antibody labeling of these thickenings indicates the presence of rhamnogalacturonan-I epitopes. Acclimatization also results in the proliferation of the cell's vacuolar networks and macroautophagy. Penium's ability to acclimatize to osmotic stress offers insight into the transition of ancient zygnematophytes from an aquatic to terrestrial existence.

Natural selection on traits and trait plasticity in Arabidopsis thaliana varies across competitive environments.

Interspecific competition reduces resource availability and can affect evolution. We quantified multivariate selection in the presence and absence of strong interspecific competition using a greenhouse experiment with 35 natural accessions of Arabidopsis thaliana. We assessed selection on nine traits representing plant phenology, growth, and architecture, as well as their plasticities. Competition reduced biomass and fitness by over 98%, and plastic responses to competition varied by genotype (significant G × E) for all traits except specific leaf area (SLA). Competitive treatments altered selection on flowering phenology and plant architecture, with significant selection on all phenology traits and most architecture traits under competition-present conditions but little indication that selection occurred in the absence of competitors. Plasticity affected fitness only in competition-present conditions, where plasticity in flowering time and early internode lengths was adaptive. The competitive environment caused changes in the trait correlation structure and surprisingly reduced phenotypic integration, which helped explain some of the observed selection patterns. Despite this overall shift in the trait correlation matrix, genotypes with delayed flowering had lower SLA (thicker, tougher leaves) regardless of the competitive environment, a pattern we have not seen previously reported in the literature. Overall, our study highlights multiple ways in which interspecific competition can alter selective regimes, contributing to our understanding of variability in selection processes over space and time.

Experimental Manipulation of Pectin Architecture in the Cell Wall of the Unicellular Charophyte, Penium Margaritaceum.

Pectins represent one of the main components of the plant primary cell wall. These polymers have critical roles in cell expansion, cell-cell adhesion and response to biotic stress. We present a comprehensive screening of pectin architecture of the unicellular streptophyte, Penium margaritaceum. Penium possesses a distinct cell wall whose outer layer consists of a lattice of pectin-rich fibers and projections. In this study, cells were exposed to a variety of physical, chemical and enzymatic treatments that directly affect the cell wall, especially the pectin lattice. Correlative analyses of pectin lattice perturbation using field emission scanning electron microscopy, confocal laser scanning microscopy, and transmission electron microscopy demonstrate that pectin lattice microarchitecture is both highly sensitive and malleable.

The Penium margaritaceum Genome: Hallmarks of the Origins of Land Plants.

The evolutionary features and molecular innovations that enabled plants to first colonize land are not well understood. Here, insights are provided through our report of the genome sequence of the unicellular alga Penium margaritaceum, a member of the Zygnematophyceae, the sister lineage to land plants. The genome has a high proportion of repeat sequences that are associated with massive segmental gene duplications, likely facilitating neofunctionalization. Compared with representatives of earlier diverging algal lineages, P. margaritaceum has expanded repertoires of gene families, signaling networks, and adaptive responses that highlight the evolutionary trajectory toward terrestrialization. These encompass a broad range of physiological processes and protective cellular features, such as flavonoid compounds and large families of modifying enzymes involved in cell wall biosynthesis, assembly, and remodeling. Transcriptome profiling further elucidated adaptations, responses, and selective pressures associated with the semi-terrestrial ecosystems of P. margaritaceum, where a simple body plan would be an advantage.

Endomembrane architecture and dynamics during secretion of the extracellular matrix of the unicellular charophyte, Penium margaritaceum.

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.

Arabinogalactan Proteins and the Extracellular Matrix of Charophytes: A Sticky Business.

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.

Genomic Admixture Between Locally Adapted Populations of Arabidopsis thaliana (mouse ear cress): Evidence of Optimal Genetic Outcrossing Distance.

Admixture can break up divergent genetic architectures between populations, resulting in phenotypic novelty and generating raw material for environmental selection. The contribution of admixture to progeny trait variation and fitness varies based on the degree of genetic isolation between the parental populations, for which most studies have used geographic distance as a proxy. A novel approach is to estimate optimal crossing distance using the adaptive genetic distance between mates estimated from loci that contribute directly to local adaptation. Here, we aim to understand the effect of admixture on disrupting local adaptation of ecotypes of Arabidopsis thaliana separated along gradients of geographic, background, and locally adaptive genetic distances. We created experimental F1 hybrids between ecotypes that vary in geographic distance and used SNP data to estimate background (putatively neutral) and adaptive genetic distance. Hybrids were grown under controlled conditions, and fitness, growth, and phenology traits were measured. The different traits measured showed a clear effect of adaptive genetic distance, but not geographic distance. The earliest bolting hybrids were intermediate in the adaptive genetic distance between their parents, and also had higher biomass and fitness in terms of fruit and seed production. Our results suggest that disruption of locally adaptive genomic loci decreases the performance of offspring between distantly related parents, but that crosses between very closely related parents also reduce performance, likely through the expression of deleterious recessive alleles. We conclude that during admixture, selection may have to balance the consequences of disrupting local adaption while also avoiding inbreeding depression.

The ubiquity of phenotypic plasticity in plants: a synthesis.

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.

PLASTICIDAD FENOTÍPICA EN Lippia alba (VERBENACEAE) EN RESPUESTA A LA DISPONIBILIDAD HÍDRICA EN DOS AMBIENTES LUMÍNICOS / Phenotypic Plasticity in Lippia alba (Verbenaceae) in Response to Water Availability in Two Light Environments

La plasticidad fenotípica es uno de los medios por los cuales las plantas pueden ajustar su morfología y fisiología, permitiéndoles enfrentar la heterogeneidad ambiental bajo condiciones naturales. El presente estudio evaluó la plasticidad fenotípica (PF) en respuesta a la disponibilidad hídrica del suelo, bajo dos condiciones de luz, en clones de dos poblaciones de Lippia alba (Verbanaceae) contrastantes en la heterogeneidad del régimen de precipitación en su hábitat natural, a través de normas de reacción morfológicas y de asignación de biomasa. El experimento se desarrolló en condiciones semicontroladas en invernadero, siguiendo un diseño experimental completamente aleatorizado, bajo un esquema factorial dos * dos (disponibilidad hídrica y poblaciones). La mayoría de los caracteres evaluados mostraron PF. No obstante, algunos caracteres de la población proveniente de condiciones naturales más homogéneas presentaron una reducción de plasticidad marcada. Lo anterior sugiere una posible relación entre el amplio rango de distribución de Lippia alba, PF y eventos de adaptación local.

The phenotypic plasticity is one possible way for plants to adjust their morphology and physiology to cope with the environmental heterogeneity of their natural conditions. This study tested the phenotypic plasticity (PF) in response to soil water availability, under two light conditions, in clones of two populations of Lippia alba (Verbanaceae), which show differences in the precipitation heterogeneity, using morphological and biomass allocation reaction norms. The experiment was conducted in a greenhouse under semi-controlled conditions in a random experimental design, under a factorial design of two * two (water availability and populations). The majority of the traits showed PF. Although, some traits of the populations that come from more homogenous conditions showed a significant reduction in plasticity. This could suggest a relationship between the wide ecological breadth of Lippia alba, PF and local adaptation events.