Parent Ramets Research Articles

Changes induced by parental neighboring touch in the clonal plant Glechoma longituba depend on the light environment.

Touch by neighboring plants is a common but overlooked environmental variable for plants, especially in dense vegetation. In addition, shade is inevitable for understory plants. The growth performance of clonal plant to the interaction between thigmomorphogenesis and shade response, and their impact on light adaptability is still unknown. At the present study, parental ramets of Glechoma longituba were exposed to two conditions (neighboring touch and shade), and their offspring ramets were in ambient or shaded environment. The phenotype and growth of parental and offspring ramets were analyzed. The results showed that neighboring touch of parental ramets regulated the performance of offspring ramets, while the effect depended on the light environment. The parental neighboring touch occurring in ambient environment suppressed the expansion of leaf organ, showed as a shorter petiole and smaller leaf area. Moreover, G. longituba exhibited both shade avoidance and shade tolerance characters to shaded environment, such as increased leaf area ratio and leaf mass ratio, longer specific petiole length and specific stolon length. It was notable that these characters of shade response could be promoted by parental neighboring touch to some extent. Additionally, parental light environment plays an important role in offspring growth, parent with ambient light always had well-grown offspring whatever the light condition of offspring, but the growth of offspring whose parent in shaded environment was inhibited. Finally, for the offspring with shaded environment, the touch between parental ramets in shade environment showed a disadvantage on their growth, but the influence of the touch between parental ramets in ambient environment was slight. Overall, the interaction of parental neighboring touch and shade environment complicate the growth of understory plants, the performance of plants is the integrated effect of both. These findings are conducive to an in-depth understanding of the environmental adaptation of plants.

Nitrogen fertilization practices alter microbial communities driven by clonal integration in Moso bamboo

Nitrogen (N) fertilization is crucial for maintaining plant productivity. Clonal plants can share resources between connected ramets through clonal integration influencing microbial communities and regulating soil biogeochemical cycling, especially in the rhizosphere. However, the effect of various N fertilization practices on microbial communities in the rhizosphere of clonal ramets remain unknown. In this study, clonal fragments of Moso bamboo (Phyllostachys edulis), consisting of a parent ramet, an offspring ramet, and an interconnecting rhizome, were established in the field. NH4NO3 solution was applied to the parent, offspring ramets or rhizomes to investigate the effect of fertilization practices on the structure and function of rhizosphere microbial communities. The differences in N availability, microbial biomass and community composition, and abundance of nitrifying genes among rhizosphere soils of ramets gradually decreased during the rapid growth of Moso bamboo, irrespective of fertilization practice. The soil N availability variation, particularly in NO3−, caused by fertilization practices altered the rhizosphere microbial community. Soil N availability and stable microbial biomass N in parent fertilization were the highest, being 9.0 % and 18.7 %, as well as 60.8 % and 90.4 % higher than rhizome and offspring fertilizations, respectively. The microbial network nodes and links in rhizome fertilization were 1.8 and 7.5 times higher than in parent and offspring fertilization, respectively. However, the diversity of bacterial community and abundance of nitrifying and denitrifying genes were the highest in offspring fertilization among three practices, which may be associated with increased N loss. Collectively, the rhizosphere microbial community characteristics depended on fertilization practices by altering the clonal integration of N in Moso bamboo. Parent and rhizome fertilization were favorable for N retention in plant-soil system and resulted in more stable microbial functions than offspring fertilization. Our findings provide new insights into precision fertilization for the sustainable Moso bamboo forest management.

DNA methylation variation and growth in the clonal Duchesnea indica is regulated by both past and present lead environments

ABSTRACT Studies suggest that clonal plants’ ability to select habitats and forage in a heterogeneous environment is influenced by their past environment, with stress legacy potentially playing a crucial role. In this study, we examined parental ramets of Duchesnea indica Focke that were subject to either a control or lead-contaminated environment (past environment), and their newborn offspring were then transplanted into control, homogeneous lead or heterogeneous lead environment (present environment). We analysed how past and present environments affect plant growth and DNA methylation in offspring. The result shown that the DNA methylation loci composition of offspring was affected by the interaction of parental environment and offspring environment, and DNA methylation levels were higher in heterogeneous environments. Moreover, our findings indicate that offspring would thrive in the heterogeneous lead environment if they did not experience lead pollution in the past, their progeny will avoid lead toxicity by reducing underground biomass allocation. However, when the parents experienced lead stress environment, their biomass allocation strategies disappeared, and they prefer to grow in favourable patches to avoid lead-contaminated patches. We concluded that the integration of historical parental exposure to lead-contaminated and current information about their offspring’s environment are impacting plant phenotypes. It is possible that the stress legacy from the parents has been transmitted to their offspring ramets, and the stress legacy is at least partly based on heritable epigenetic variation. The phenotypic variation regulated by the stress legacy affects the growth performance, biomass allocation strategy, and even the behaviour of D. indica.

Reproductive Responses to Increased Shoot Density and Global Change Drivers in a Widespread Clonal Wetland Species, Schoenoplectus americanus

The expansion of many wetland species is a function of both clonal propagation and sexual reproduction. The production of ramets through clonal propagation enables plants to move and occupy space near parent ramets, while seeds produced by sexual reproduction enable species to disperse and colonize open or disturbed sites both near and far from parents. The balance between clonal propagation and sexual reproduction is known to vary with plant density but few studies have focused on reproductive allocation with density changes in response to global climate change. Schoenoplectus americanus is a widespread clonal wetland species in North America and a dominant species in Chesapeake Bay brackish tidal wetlands. Long-term experiments on responses of S. americanus to global change provided the opportunity to compare the two modes of propagation under different treatments. Seed production increased with increasing shoot density, supporting the hypothesis that factors causing increased clonal reproduction (e.g., higher shoot density) stimulate sexual reproduction and dispersal of genets. The increase in allocation to sexual reproduction was mainly the result of an increase in the number of ramets that flowered and not an increase in the number of seeds per reproductive shoot, or the ratio between the number of flowers produced per inflorescence and the number of flowers that developed into seeds. Seed production increased in response to increasing temperatures and decreased or did not change in response to increased CO2 or nitrogen. Results from this comparative study demonstrate that plant responses to global change treatments affect resource allocation and can alter the ability of species to produce seeds.

Predictability of parental ultraviolet-B environment shapes the growth strategies of clonal Glechoma longituba

Although there is an increasing debate about ecological consequences of environmental predictability for plant phenotype and fitness, the effect of predictability of parental environments on the offspring is still indefinite. To clarify the role of environmental predictability in maternal effects and the growth strategy of clonal offspring, a greenhouse experiment was conducted with Glechoma longituba. The parental ramets were arranged in three ultraviolet-B (UV-B) conditions, representing two predictable environments (regular and enhanced UV-B) and an unpredictable environment (random UV-B), respectively. The offspring environments were the same as their parent or not (without UV-B). At the end of experiment, the growth parameters of offspring were analyzed. The results showed that maternal effects and offspring growth were regulated by environmental predictability. Offspring of unpredictable environmental parents invested more resources in improving defense components rather than in rapid growth. Although offspring of predictable parents combined two processes of defense and growth, there were still some differences in the strategies between the two offspring, and the offspring of regular parent increased the biomass allocation to roots (0.069 g of control vs. 0.092 g of regular), but that of enhanced parent changed the resource allocation of nitrogen in roots and phosphorus in blade. Moreover, when UV-B environments of parent and offspring were matched, it seemed that maternal effects were not adaptive, while the growth inhibition in the predictable environment was weaker than that in unpredictable environment. In the predictable environment, the recovered R/S and the increased defense substances (flavonoid and anthocyanin) contributed to improving offspring fitness. In addition, when UV-B environments of parent and offspring were mismatched, offspring growth was restored or improved to some extent. The offspring performance in mismatched environments was controlled by both transgenerational effect and within-generational plasticity. In summary, the maternal effects affected growth strategies of offspring, and the differences of strategies depended on the predictability of parental UV-B environments, the clone improved chemical defense to cope with unpredictable environments, while the growth and defense could be balanced in predictable environments. The anticipatory maternal effects were likely to improve the UV-B resistance.

Clonal Parental Effects on Offspring Growth of Different Vegetative Generations in the Aquatic Plant Pistia stratiotes.

Parental (environmental) effects can modify the growth of offspring, which may play an essential role in their adaptation to environmental variation. While numerous studies have tested parental effects on offspring growth, most have considered offspring growth of only one generation and very few have considered offspring growth of different generations. We conducted a greenhouse experiment with an aquatic clonal plant Pistia stratiotes. We grew a single ramet of P. stratiotes under low or high nutrients, the initial (parent) ramets produced three different generations of offspring ramets, and these offspring ramets were also subjected to the same two nutrient levels. High nutrients currently experienced by the offspring increased biomass accumulation and ramet number of all three offspring generations of P. stratiotes. However, these positive effects on biomass were greater when the offspring ramets originated from the parent ramets grown under low nutrients than when they were produced by the parent ramets grown under high nutrients. These results suggest that parental effects can impact the performance of different offspring generations of clonal plants. However, heavier offspring ramets produced under high nutrients in parental conditions did not increase the subsequent growth of the offspring generations. This finding indicates that parental provisioning in favorable conditions may not always increase offspring growth, partly depending on root allocation but not ramet size such as ramet biomass.

Genotypic differences in response to different patterns of clonal fragmentation in the aquatic macrophytePistia stratiotes

AbstractConnected individuals (ramets) of clonal plants are frequently fragmented due to disturbance, and such clonal fragmentation may influence their growth. However, it is unclear whether different patterns of clonal fragmentation produce differential effects on plant growth and whether such differences vary with genotypes. We collected one group of connected ramets of the stoloniferous floating invasive macrophyte Pistia stratiotes from each of six provinces (Guangdong, Guangxi, Hubei, Jiangsu, Sichuan and Yunnan Provinces) of China, and assumed that these ramets belonged to different genotypes. After several generations of vegetative propagation to remove potential parental effects, new ramets (hereafter referred to as parent ramets) from each of these assumed genotypes were subjected to four patterns of clonal fragmentation, i.e., all offspring ramets produced during the experiment remained connected to their parent ramet (control), or the primary (1°), secondary (2°) or tertiary (3°) offspring ramets were disconnected from their parent ramet. We found significant growth differences among the assumed genotypes, confirming that the ramets sampled from these six provinces were from different genotypes. Compared with the control, disconnecting the 1°, 2° or 3° ramets did not significantly affect biomass of the whole clone (i.e., the parent ramet plus all offspring ramets) of P. stratiotes. Disconnecting the 1° ramets significantly increased biomass of the parental ramet of all six genotypes compared with the control. However, disconnecting the 2° or 3° ramets had little effect on biomass of the parent ramet for all genotypes except the Guangxi genotype, for which disconnecting the 2° ramets decreased biomass of the parent ramet. Disconnecting the 1°, 2° or 3° ramets had no impact on biomass or number of the 1° or the 3° ramets for any of the genotypes. However, disconnecting the 1° ramets decreased biomass of the 2° ramets for the Guangdong and Yunnan genotypes, but had no effect for the other four genotypes. We conclude that different patterns of clonal fragmentation can have different impacts on parent and offspring ramets of P. stratiotes and such an impact may vary with genotypes.

Parental UV-B radiation regulates the habitat selection of clonal Duchesnea indica in heterogeneous light environments.

Habitat selection behaviour is an effective strategy adopted by clonal plants in heterogeneous understorey light environments, and it is likely regulated by the parental environment's ultraviolet-B radiation levels (UV-B) due to the photomorphogenesis of UV-B and maternal effects. Here, parental ramets of Duchesnea indica were treated with two UV-B radiation levels [high (UV5 group) and low (UV10 group)], newborn offspring were grown under a heterogeneous light environment (ambient light vs shade habitat), and the growth and DNA methylation variations of parents and offspring were analysed. The results showed that parental UV-B affected not only the growth of the parent but also the offspring. The offspring of different UV-B-radiated parents showed different performances. Although these offspring all displayed a tendency to escape from light environments, such as entering shade habitats earlier, and allocating more biomass under shade (33.06% of control, 42.28% of UV5 and 72.73% of UV10), these were particularly obvious in offspring of the high UV-B parent. Improvements in epigenetic diversity (4.77 of control vs 4.83 of UV10) and total DNA methylation levels (25.94% of control vs 27.15% of UV10) and the inhibition of shade avoidance syndrome (denser growth with shorter stolons and internodes) were only observed in offspring of high UV-B parents. This difference was related to the eustress and stress effects of low and high UV-B, respectively. Overall, the behaviour of D. indica under heterogeneous light conditions was regulated by the parental UV-B exposure. Moreover, certain performance improvements helped offspring pre-regulate growth to cope with future environments and were probably associated with the effects of maternal DNA methylation variations in UV-B-radiated parents.

Ultraviolet B Radiation Triggers DNA Methylation Change and Affects Foraging Behavior of the Clonal Plant Glechoma longituba.

Clonal plants in heterogeneous environments can benefit from their habitat selection behavior, which enables them to utilize patchily distributed resources efficiently. It has been shown that such behavior can be strongly influenced by their memories on past environmental interactions. Epigenetic variation such as DNA methylation was proposed to be one of the mechanisms involved in the memory. Here, we explored whether the experience with Ultraviolet B (UV-B) radiation triggers epigenetic memory and affects clonal plants’ foraging behavior in an UV-B heterogeneous environment. Parental ramets of Glechoma longituba were exposed to UV-B radiation for 15 days or not (controls), and their offspring ramets were allowed to choose light environment enriched with UV-B or not (the species is monopodial and can only choose one environment). Sizes and epigenetic profiles (based on methylation-sensitive amplification polymorphism analysis) of parental and offspring plants from different environments were also analyzed. Parental ramets that have been exposed to UV-B radiation were smaller than ramets from control environment and produced less and smaller offspring ramets. Offspring ramets were placed more often into the control light environment (88.46% ramets) than to the UV-B light environment (11.54% ramets) when parental ramets were exposed to UV-B radiation, which is a manifestation of “escape strategy.” Offspring of control parental ramets show similar preference to the two light environments. Parental ramets exposed to UV-B had lower levels of overall DNA methylation and had different epigenetic profiles than control parental ramets. The methylation of UV-B-stressed parental ramets was maintained among their offspring ramets, although the epigenetic differentiation was reduced after several asexual generations. The parental experience with the UV-B radiation strongly influenced foraging behavior. The memory on the previous environmental interaction enables clonal plants to better interact with a heterogeneous environment and the memory is at least partly based on heritable epigenetic variation.

Epigenetic memory and growth responses of the clonal plant Glechoma longituba to parental recurrent UV-B stress.

The responses of plants to recurrent stress may differ from their responses to a single stress event. In this study, we investigated whether clonal plants can remember past environments. Parental ramets of Glechoma longituba (Nakai) Kuprian were exposed to UV-B stress treatments either once or repeatedly (20 and 40 repetitions). Differences in DNA methylation levels and growth parameters among parents, offspring ramets and genets were analysed. Our results showed that UV-B stress reduced the DNA methylation level of parental ramets, and the reduction was enhanced by increasing the number of UV-B treatments. The epigenetic variation exhibited by recurrently stressed parents was maintained for a long time, but that of singly stressed parents was only short-term. Moreover, clonal plants responded to different UV-B stress treatments with different growth strategies. The one-time stress was a eustress that increased genet biomass by increasing offspring leaf allocation and defensive allocation in comparison to the older offspring. In contrast, recurring stress was a distress that reduced genet biomass, increased the biomass of storage stolons, and allocated more defensive substances to the younger ramets. This study demonstrated that the growth of offspring and genets was clearly affected by parental experience, and parental epigenetic memory and the transgenerational effect may play important roles in this effect.

With a little help from my friends: physiological integration facilitates invasion of wetland grass Elymus athericus into flooded soils

Tidal wetlands worldwide are undergoing rapid invasions by tall‐growing clonal grasses. Prominent examples are invasions by species of the genera Spartina, Phragmites and Elymus. The responsible physiological and ecological drivers of these invasions are poorly understood. Physiological integration (PI) is a key trait of clonal plants, which enables the exchange of resources among ramets. We investigated PI in Elymus athericus, which has been rapidly spreading from high‐marsh into low‐marsh environments of European salt marshes during the last decades. We applied a nitrogen stable‐isotope approach to trace nutrient translocation between ramets in a factorial mesocosm experiment. The experiment was set up to mimic an invasion pattern commonly found in tidal wetlands, i.e. from high‐elevated and rarely flooded into low‐elevated and frequently flooded microenvironments. We tested for intraspecific variability in PI by including two genotypes of Elymus that naturally occur at different elevations within the tidal frame, a high‐marsh (HM) and a low‐marsh (LM) genotype. PI strongly increased offspring ramet aboveground and belowground biomass by 62 and 81%, respectively. Offspring ramets under drained conditions had 95% greater belowground biomass than those under flooded conditions. LM genotype offspring ramets produced 27% more aboveground biomass than HM genotypes. Offspring ramets were clearly more enriched in 15N under flooded versus drained conditions; however, this positive effect of flooding on δ15N was only significant in the LM genotype. Our findings demonstrate the importance of PI for the growth of Elymus offspring ramets and thereby for the species' capacity for fast vegetative spread. We show that offspring ramets under stressful flooded conditions are more dependent on nutrient supply from parent ramets than those under drained conditions. Our data furthermore suggest a higher degree of adaptation to flooding via PI in the LM versus HM genotype. In conclusion, we highlight the importance of assessing PI and intraspecific trait variability to understand invasion processes within ecosystems.

Nitrogen deposition regulates the clonal growth of Leymus chinensis, a typical clonal plant in arid and semi-arid regions

Purpose The purpose of this study is to examine the effects of nitrogen (N) deposition on clonal growth in a rhizome clonal plant, Leymus chinensis (Trin.) Tzvel. Design/methodology/approach The study established seven N concentration gradients (0, 2, 4, 8, 16, 32 and 64 g N m−2) to simulate the continuous increase in N deposition for the cultivation of L. chinensis seedlings and assess the response mechanism of the cloned L. chinensis plant at different N levels by analyzing the aboveground and belowground plant appearance traits, parent ramets and daughter ramets of resource allocation and biomass allocation. Findings The results of this study showed that the different N treatment levels could promote clonal growth and had certain regularity under the seven treatments. The addition of N could significantly increase the ramet number, rhizome length, rhizome spacer length, biomass of mother ramets, daughter ramets and belowground L. chinensis population when the N addition was greater than 4 g m−2; however, the clonal growth ability of L. chinensis decreased and the rhizome length, ramet number, stem and leaf biomass of daughter ramets and stem biomass of mother ramets significantly decreased when the N addition was greater than 32 g N m−2. Originality/value With global warming, atmospheric N deposition is increasing and it is of great significance to explore the response mechanism of different N levels for the growth of clone plants. This study provides basic data and a theoretical basis for the survival prediction of cloned plants under the background of a global climate change strategy and has important theoretical and practical significance for the scientific management of grasslands in the future.

Effects of fragmentation of clones compound over vegetative generations in the floating plant Pistia stratiotes.

Clonal plants dominate many plant communities, especially in aquatic systems, and clonality appears to promote invasiveness and to affect how diversity changes in response to disturbance and resource availability. Understanding how the special physiological and morphological properties of clonal growth lead to these ecological effects depends upon studying the long-term consequences of clonal growth properties across vegetative generations, but this has rarely been done. This study aimed to show how a key clonal property, physiological integration between connected ramets within clones, affects the response of clones to disturbance and resources in an aquatic, invasive, dominant species across multiple generations. Single, parental ramets of the floating stoloniferous plant Pistia stratiotes were grown for 3 weeks, during which they produced two or three generations of offspring; connections between new ramets were cut or left intact. Individual offspring were then used as parents in a second 3-week iteration that crossed fragmentation with previous fragmentation in the first iteration. A third iteration yielded eight treatment combinations, zero to three rounds of fragmentation at different times in the past. The experiment was run once at a high and once at a low level of nutrients. In each iteration, fragmentation increased biomass of the parental ramet, decreased biomass of the offspring and increased number of offspring. These effects persisted and compounded from one iteration to another, though more recent fragmentation had stronger effects, and were stronger at the low than at the high nutrient level. Fragmentation did not affect net accumulation of mass by groups after one iteration but increased it after two iterations at low nutrients, and after three iterations at both nutrient levels. Both the positive and negative effects of fragmentation on clonal performance can compound and persist over time and can be stronger when resource levels are lower. Even when fragmentation has no short-term net effect on clonal performance, it can have a longer-term effect. In some cases, fragmentation may increase total accumulation of mass by a clone. The results provide the first demonstration of how physiological integration in clonal plants can affect fitness across generations and suggest that increased disturbance may promote invasion of introduced clonal species via effects on integration, perhaps especially at lower nutrient levels.

Effects of parental light environment on growth and morphological responses of clonal offspring.

Environments experienced by parent ramets of clonal plants can potentially influence fitness of clonal offspring ramets. Such clonal parental effects may result from heritable epigenetic changes, such as DNA methylation, which can be removed by application of DNA de-methylation agents such as 5-azacytidine. To test whether parental shading effects occur via clonal generation and whether DNA methylation plays a role in such effects, parent plants of the clonal herb Alternanthera philoxeroides were first subjected to two levels of light intensity (high versus low) crossed with two levels of DNA de-methylation (no or with de-methylation by application of 5-azacytidine), and then clonal offspring taken from each of these four types of parent plant were subjected to the same two light levels. Parental shading effects transmitted via clonal generation decreased growth and modified morphology of clonal offspring. Offspring responses were also influenced by DNA methylation level of parent plants. For clonal offspring growing under low light, parental shading effects on growth and morphology were always negative, irrespective of the parental de-methylation treatment. For clonal offspring growing under high light, parental shading effects on offspring growth and morphology were negative when the parents were not treated with 5-azacytidine, but neutral when they were treated with 5-azacytidine. Overall, parental shading effects on clonal offspring performance of A.philoxeroides were found, and DNA methylation is likely to be involved in such effects. However, parental shading effects contributed little to the tolerance of clonal offspring to shading.

Effects of Transgenerational Plasticity on Morphological and Physiological Properties of Stoloniferous Herb Centella asiatica Subjected to High/Low Light.

Environmentally induced transgenerational plasticity can increase success of progeny and thereby be adaptive if progeny experiences the similarly parental environment. The ecological and evolutionary significance of transgenerational plasticity in plant has been studied mainly in the context of sexual generations. A pot experiment using the stoloniferous herb Centella asiatica was conducted to investigate the effects of high/low light treatment experienced by parental ramets (F0 generation) on morphological and physiological properties of offspring ramets (F2 generation) as well as growth performance. Light environment experienced by parental ramets (F0 generation) significantly influenced petiole length, specific petiole length, internode length of stolon, leaf area, specific leaf area (SLA), leaf nitrogen and chlorophyll contents, potential maximum net photosynthetic rate (Pmax) in offspring ramets subjected to parental or non-parental environments even after they were detached from the parental ramets. Potential maximum net photosynthetic rate (Pmax) of offspring ramets (F2 generation) from parental ramets (F0 generation) subjected to low light treatment was significantly greater than that of offspring ramets (F2 generation) from parental ramets (F0 generation) subjected to high light treatment. Potential maximum net photosynthetic rate (Pmax) of offspring ramets (F2 generation) subjected to parental light environment was greater than that of offspring ramets (F2 generation) subjected to non-parental light environment. The greatest biomass accumulation and total stolon length were observed in offspring ramets (F2 generation) subjected to low light treatment as parental ramets (F0 generation) experienced. When parental ramets (F0 generation) were subjected to low light treatment, biomass accumulation and total stolon length of offspring ramets (F2 generation) experiencing parental light environment were significantly greater than those of offspring ramets (F2 generation) experiencing non-parental light environment. Opposite pattern was observed in offspring ramets (F2 generation) from parental ramets subjected to high light treatment. Our work provides evidence that transgenerational plasticity through both morphological and physiological flexibility was triggered across vegetative generations for stoloniferous herb C. asiatica subjected to high/low light treatment. The transgenerational plasticity can allow offspring ramets to present adaptive phenotype early without lag time in response to the current environment. Thus, it is very important for clonal plants in adapting temporally and spatially heterogeneous habitats.

Physiological integration of growth, carbohydrates, and soluble protein of Zoysia japonica clonal ramets under nutrient heterogeneity

This study was carried out to analyze the changes of growth, the contents of carbohydrates and soluble protein of Zoysia japonica clonal ramets under nutrient heterogeneity, where the connected and disconnected ramets were treated with different nutrient levels. The results indicated that under the nutrient heterogeneity the parent ramets in middle or high nutrient levels improved the aboveground biomass, belowground biomass, and total biomass, with the enhancement of 32.5%, 22.1% and 24.8% at high nutrient level, respectively, reduced the root/shoot ratio, the contents of soluble sugar and non-structural carbohydrates (NSC), with the reduction of 7.7%, 15.2% and 13.1% in high nutrient level, respectively, but had no significant impacts on the contents of starch, cellulous, and soluble protein of the connected daughter ramets. The daughter ramets in middle and high nutrient levels had no significant impacts on the growth and the contents of carbohydrates and soluble protein of the connected parent ramets. There was a significant physiological integration effects from parent to the daughter ramets on the biomass, root/shoot, the contents of soluble sugar and NSC. The intensity of physiological integration was proportional to the nutrient gradient of ramets, but had no significant physiological integration effects on the contents of starch, cellulous and soluble protein. The daughter ramets had no physiological integration for any indicator of the parent ramets. There was a unidirectional physiological integration between parent and daughter ramets of Z. japonica.

Clonal plant Duchesnea indica Focke forms an effective survival strategy in different degrees of Pb-contaminated environments

Clonal plants in heterogeneous environments can benefit from habitat selection, giving them the ability to utilize patchily distributed resources efficiently. However, most research is conducted in a resource heterogeneous environment, and the study of heterogeneous environment of non-resource material (copper, cadmium, lead, and so on) is limited. Research into clonal plant growth under heterogeneous toxic conditions could contribute to our understanding of the strategy of the selective establishment pattern. Thus, we examined clonal growth in a heterogeneous lead environment to enhance understanding of habitat selection strategies. The growth indices (stolon-length, ramet number, biomass, and lead-concentrations) of Duchesnea indica were examined under three levels (low, moderate, and high were represented by 0, 50 and 100%) of lead contamination and two degrees of heterogeneity (low and high heterogeneity under moderate contamination) environments in a glasshouse study. Habitat selection strategy was analyzed by clone growth pattern, labor division, and risk-spreading. The clones under the moderate contamination level, especially with high heterogeneity, demonstrated the optimal growth. They expanded their growth-pattern to escape the toxic environment, and rooted more ramets in the unpolluted patches, allocating more aboveground biomass to these areas. Moreover, parent ramets transported their lead in the soil to the offspring. The offspring spread the toxic risk by accumulating lead in their roots and producing more ramets. Optimal growth of clonal plants occurred in environments moderately contaminated with lead, especially under higher heterogeneity, which performed effective survival strategy by expensive growth architecture, aboveground biomass increase and risk-spreading.

Physiological integration of growth and photosynthesis of Zoysia japonica clonal ramets under nutrient heterogeneity

This study was carried out to analyze the changes of growth and photosynthesis of clonal ramets under nutrient heterogeneity, where the connected and disconnected ramets were treated with different nutrient levels. The results showed that under the nutrient heterogeneity the parent ramets in middle or high nutrient levels improved leaf length, leaf width, root mass, leaf mass, net photosynthetic rate, stomatal conductance, transpiration rate and water use efficiency of the connected daughter ramets, with an increase of 16.0%, 8.3%, 24.4%, 58.1%, 30.3%, 54.0%, 9.2% and 21.9% in high nutrient level, respectively, but reduced the root/shoot and intercellular CO2 concentration of the connected daughter ramets, with a decreases of 21.6% and 31.5% in high nutrient level, respectively. In contrast, the daughter ramets in the middle or high nutrient level had no significant impacts on the growth and photosynthesis of the connected parent ramets. There was a physiological integration from the parent ramets to the daughter ramets. The larger the nutrient differences of ramets was, the stronger the intensity of physiological integration was. The daughter ramets were the unidirectional beneficiary from the physiological integration, as the daughter ramets benefited from the parent ramets but had no positive effects on the daughter ramets.

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.

Clonal integration in Ludwigia hexapetala under different light regimes

Physiological integration among ramets of invasive plant species may support their colonization and spread in novel aquatic environments where growth-limiting resources are spatially heterogeneous. Under contrasting light conditions, we investigated how clonal integration influences growth, biomass allocation and morphology of Ludwigia hexapetala, an emergent floating-leaved macrophyte that is highly invasive in a range of wetland habitat types. In aquatic mesocosms, stolons of offspring ramets were either connected or severed from parent plants, with the pairs exposed to homogenous or heterogeneous combinations of sun or 85% shade. Morphological traits of all ramets were strongly influenced by light environment, and low light availability decreased plant growth, regardless of integration status. Allocation patterns varied with light regime; shaded plants increased allocation to leaf biomass while sun plants allocated more resources to belowground growth. Offspring ramets integrated with parents produced more biomass, suggesting a fitness advantage through integration. However, parent ramet performance declined with stoloniferous integration; integrated parents produced fewer ramets and allocated more resources to belowground biomass. For most response variables measured, there was no significant interactive effect between light treatment and integration, although parents growing in the shade attached to an offspring in the sun increased root mass ratio. The ability to establish and spread into new environments is a key trait of invasive plants, and physiological integration of resources may improve the establishment of juvenile ramets across variable light environments during early colonization. Physiological integration in patchy light environments may contribute to the invasiveness of L. hexapetala.