Effects of Glyphosate Application on Physiologically Integrated Clones of the Invasive Plant Carpobrotus edulis

Management of invasive alien species is a high priority for biodiversity conservation. Here, we studied the effects of glyphosate application, at 0.06 g/m2 concentration, on physiologically integrated basal and apical ramets of the invasive clonal plant Carpobrotus edulis. Physiological integration allows the transport of resources and other substances between connected ramets in clonal plants. We found a significant reduction of growth and photochemical efficiency both in basal and apical ramets of C. edulis after glyphosate application. Interestingly, we also observed a significant growth reduction in untreated basal ramets when they remained connected to apical ramets treated with glyphosate. This result was interpreted as a cost for basal ramets due to supporting severely stressed apical ramets. Therefore, local application of glyphosate to apical ramets of C. edulis can negatively affect not only their own growth, but also the growth of their interconnected, untreated basal ramets. Our results suggest that glyphosate effectiveness can be maintained when applied only to one part of the clone so that the amount of herbicide used in eradication programs can be greatly reduced, which can minimize the negative impact of chemical herbicides on ecosystems.

Changes in the growth and reproduction of a clonal plant as a result of disruption of mycorrhizal network

In a clonal network, a mother plant is connected with daughter ramets. During network development, new ramets may encounter barriers that disrupt network integrity. As a result, resource allocation within a network is disturbed. In this study, the effect of network integrity disruption on the size of ramets and their sexual reproduction was investigated in mouse-ear hawkweed (Hieracium pilosella). Three types of networks were formed experimentally with unlimited resource allocation, with limited resource allocation between a mother plant and its daughter ramets and with limited resource allocation between all ramets. Networks were either supported by the presence of a mycorrhizal fungus or restricted by its absence. We found that the size of the mother and the effectiveness of sexual reproduction did not differ among network types. The length and dry mass of runners were higher in cases with limited resource exchange between a mother plant and its daughters. In the clonal plant network without any barriers to connection, a higher number of rosettes and lower dry mass of daughters were recorded. The mean number of daughter flowers did not differ among the network types. Mycorrhizal network is one of the most important factors for the sexual reproduction of clonal plants. With a reduced mycorrhizal network, plants invested in clonal growth.

Time dynamics of stress legacy in clonal transgenerational effects: a case study on Trifolium repens

Stress can be remembered by plants in a form of stress legacy that can alter future phenotypes of previously stressed plants and even phenotypes of their offspring. DNA methylation belongs among the mechanisms mediating the stress legacy. It is however not known for how long the stress legacy is carried by plants. If the legacy is long lasting, it can become maladaptive in situations when parental-offspring environments do not match. We investigated for how long after the last exposure of a parental plant to drought can the phenotype of its clonal offspring be altered. We grew parental plants of three genotypes of Trifolium repens for five months either in control conditions or in control conditions that were interrupted with intense drought periods applied for two months in four different time-slots. We also treated half of the parental plants with a demethylating agent (5-azaC) to test for the potential role of DNA methylation in the stress legacy. Then, we transplanted parental cuttings (ramets) individually to control environment and allowed them to produce offspring ramets for two months. The drought stress experienced by parents affected phenotypes of offspring ramets. The stress legacy resulted in enhanced number of offspring ramets originating from parents that experienced drought stress even 8 weeks before their transplantation to the control environment. 5-azaC altered transgenerational effects on offspring ramets. We confirmed that drought stress can trigger transgenerational effect in T. repens that is very likely mediated by DNA methylation. Most importantly, the stress legacy in parental plants persisted for at least 8 weeks suggesting that the stress legacy can persist in a clonal plant Trifolium repens for relatively long period. We suggest that the stress legacy should be considered in future ecological studies on clonal plants.

Distinct responses of frond and root to increasing nutrient availability in a floating clonal plant

Current knowledge on responses of aquatic clonal plants to resource availability is largely based on studies manipulating limited resource levels, which may have failed to capture the “big picture” for aquatic clonal plants in response to resource availability. In a greenhouse experiment, we grew the floating clonal plant Spirodela polyrhiza under ten nutrient levels (i.e., 1/64×, 1/32×, 1/16×, 1/8×, 1/4×, 1/2×, 1×, 2×, 4× and 8×full-strength Hoagland solution) and examined their responses in terms of clonal growth, morphology and biomass allocations. The responses of total biomass and number of ramets to nutrient availability were unimodal. A similar pattern was found for frond mass, frond length and frond width, even though area per frond and specific frond area fluctuated greatly in response to nutrient availability. In contrast, the responses of root mass and root length to nutrient availability were U-shaped. Moreover, S. polyrhiza invested more to roots under lower nutrient concentrations. These results suggest that nutrient availability may have distinct influences on roots and fronds of the aquatic clonal plant S. polyrhiza, resulting in a great influence on the whole S. polyrhiza population.

Genotype, Environment, Year, and Harvest Effects on Fruit Quality Traits of Five Blueberry (Vaccinium corymbosum L.) Cultivars

Blueberries (Vaccinium spp.) comprise a broad range of perennial woody species. Introgression of native species into cultivated germplasm has adapted Vaccinium germplasm to a range of climates and growing conditions for cultivated blueberry. Genetic differences signify phenotypic variance that is observed among blueberry accessions. In addition, variability in geographic and climatic growing conditions between environments or within the same environment across different years may further affect fruit and plant phenotypic expression. As a result, a phenotype is a function of genetic background (G), environment (E), and their interaction (G × E). In addition, other temporally regulated factors such as year (Y) and harvest time (H) impact plant and fruit quality phenotypic variation. Our research aimed to assess the genotypic performance of five blueberry cultivars, including ‘Echota’, ‘O’Neal’, ‘Reveille’, ‘Summit’, and ‘Sunrise’. The selected cultivars were phenotyped for various fruit quality-related traits over two sequential harvests in two years and two locations. Our results indicated that genotype was a significant source of variation for most phenotypic characteristics. Further, the effect of Y × H and G × Y × H significantly affected the majority of studied phenotypic traits. Within the studied genotypes, ‘Reveille’ and ‘O’Neal’ phenotypic stability were consistent across locations and years; additionally, ‘Summit’ phenotypic characteristics were stable across years, environments, and harvests. Clonal plant replicates within a genotype, harvest, and environment, in addition to individual fruit measures, were the most significant sources of variability.

Effect of Environmental Stress on the Nutrient Stoichiometry of the Clonal Plant Phragmites australis in Inland Riparian Wetlands of Northwest China

Clonal plants play an important role in determining ecosystem properties such as community stability, species diversity and nutrient cycling. However, relatively little information is available about the stoichiometric characteristics of clonal plants and their drivers in inland riparian wetlands under strong environmental stress. In this manuscript, we studied the clonal plant Phragmites australis in an inland riparian wetland of Northwest China and compared its nutrient distribution and stoichiometry trade-offs as well as its responses to soil environmental factors in three different environments, namely, a wetland, a salt marsh, and a desert. We found that (1) P. australis could adapt to heterogeneous environments by changing its nutrient allocation strategies, as evidenced by the significant decrease in N and P concentrations, and significant increase in whole-plant C:P and N:P ratios from the wetland to the desert habitats. (2) P. australis adapted to stressful environments by changing its nutrient allocation patterns among different modules, showing a greater tendency to invest N and P in underground modules (rhizomes and roots) and an increase in the utilization efficiency of N and P in the leaves, and stems as environmental stress increased. (3) The C-N, C-P, and N:P-C in the whole plant and in each module showed significant anisotropic growth relationships in the three habitats (P < 0.05). (4) Soil water, pH and salt were the main factors limiting nutrient stoichiometry. The results of this study clarified the ecological adaptation mechanism of the clonal plant P. australis to heterogeneous environments and provided targeted protection strategies for inland riparian wetlands in Northwest China.

Transgenerational non-genetic inheritance has fitness costs and benefits under recurring stress in the clonal duckweed Spirodela polyrhiza

Although non-genetic inheritance is thought to play an important role in plant ecology and evolution, evidence for adaptive transgenerational plasticity is scarce. Here, we investigated the consequences of copper excess on offspring defences and fitness under recurring stress in the duckweed Spirodela polyrhiza across multiple asexual generations . Growing large monoclonal populations (greater than 10 000 individuals) for 30 generations under copper excess had negative fitness effects after short and no fitness effect after prolonged growth under recurring stress. These time-dependent growth rates were likely influenced by environment-induced transgenerational responses, as propagating plants as single descendants for 2 to 10 generations under copper excess had positive, negative or neutral effects on offspring fitness depending on the interval between initial and recurring stress (5 to 15 generations). Fitness benefits under recurring stress were independent of flavonoid accumulations, which in turn were associated with altered plant copper concentrations. Copper excess modified offspring fitness under recurring stress in a genotype-specific manner, and increasing the interval between initial and recurring stress reversed these genotype-specific fitness effects. Taken together, these data demonstrate time- and genotype-dependent adaptive and non-adaptive transgenerational responses under recurring stress, which suggests that non-genetic inheritance alters the evolutionary trajectory of clonal plant lineages in fluctuating environments.

Remember or not? For how long can a clonal plant remember drought stress?

Stress can be remembered by plants in a form of ‘stress memory’ that can alter future phenotypes of previously stressed plants and even phenotypes of their offspring. It was shown that DNA methylation is among the mechanisms mediating the memory. It is not known for how long the memory is kept by plants. If the memory is long lasting, it can become maladaptive in situations when parental-offspring environment differ. We investigated for how long can a parental plant “remember” that it experienced a stress and pass the memory to its clonal offspring. We grew parental plants of three genotypes of Trifolium repens for five months either in control conditions or in control conditions that were interrupted with drought pulses applied for two months in four different time-slots. We also treated half of the parental plants with 5-azacytidine (5-azaC) to test for the potential role of DNA methylation in the stress memory. Then, we transplanted parental cuttings (ramets) individually to control environment and allowed them to produce offspring ramets for two months. The drought stress experienced by parents affected phenotypes of offspring ramets. Such a memory resulted in enhanced number of offspring side branches originating from plants that experienced drought stress maximally 6 weeks before their transplantation to control environment. We did not find any transgenerational memory in offspring of plants that experienced drought stress later than 6 weeks before their transplantation. 5-azaC also reduced the effect of transgenerational memory on offspring ramets. We confirmed that drought stress can trigger transgenerational memory in T. repens that is very likely mediated by DNA methylation. Most importantly, the memory was time limited and was gradually erased. We conclude that the time limited memory on environmental stress can be adaptive as climate tends to be variable and parental-offspring environmental conditions often do not match.

Evolutionary responses of a dominant plant along a successional gradient in a salt-marsh system

The ecological responses of plant populations along a successional gradient have been intensively examined; however, the evolutionary responses received much less attention. Here, I explored genetic changes of key phenotypic traits of a dominant clonal plant (Elytrigia atherica) along a saltmarsh successional gradient by collecting samples along the successional gradient in the high and low marsh and growing them in a common environment (greenhouse). Additionally, to explore whether changes in traits are driven by abiotic (e.g. clay thickness) and biotic (e.g. grazing intensity) variables along the successional gradient, I measured these two variables in the field. I found that clay thickness (a proxy of total nitrogen) increased along the successional gradient both in the high and low marsh; grazing intensity from hares (the most important herbivores) decreased along the successional gradient in the high marsh but did not change in the low marsh. Meanwhile, I found that growth in number of leaves and ramets decreased, while rhizome length increased, along the successional gradient for E. atherica collected from the high marsh. Opposite trends were found for E. atherica collected from the low marsh. Results suggest that, in the high marsh, herbivores may overrule nutrients to drive trait changes. That is, at the early successional stages, E. atherica had higher growth in number of leaves and ramets to compensate for high-intensity grazing. In the low marsh, nutrients may be the dominant driver for trait changes. That is, at the late successional stages, E. atherica had higher growth in number of leaves and ramets but shorter rhizomes to maximize its expansion under the favorable conditions (higher nutrient availability). Results suggest that ecologically important abiotic and biotic variables such as nutrients and herbivores may also have a substantial evolutionary impact on plant populations.

Frequency of flower visitors and achene production increase with rising population size in the self-incompatible herb Centaurea scabiosa (Asteraceae)

Self-incompatible, non-clonal plant species are especially dependent on the activity of flower visitors for seed production. Therefore, populations of these plants are vulnerable to a reduced flower visitation rates, but also to increased isolation by extinction of local populations. To study how local populations of Centaurea scabiosa, a self-incompatible, bee-pollinated herb species changed over time in the region of Bonn, we collected historical population records in the area and investigated their current status. We found that more than half of the subpopulations mentioned in the literature between 1950 and 2012 (38 of 65) have since disappeared. Small populations were most vulnerable, whereas medium to large populations increased in size. In a second step, we studied visitation frequencies and achene production and weight across 14 extant populations. We found that both flower visitation frequency and achene production were positively related to population size. Achene weight was neither related to the frequency of flower visitors nor to achene production. These results clearly indicate that reduced pollen transfer and lower pollination rates may contribute to local extinction in small populations of Centaurea scabiosa. Overall our results call for an intensified monitoring of the populations of self-incompatible plant species.