Plant-soil Feedback Research Articles

Asymmetric sharing of generalist pathogens between exotic and native plants correlates with exotic impact in communities

Abstract As exotic plants invade into a new range, they can escape from specialist enemies. However, they may support generalist enemies, including both native and introduced fungal pathogens, which creates the potential for spillover and apparent competition from exotic to native plants in communities. To assess the potential for spillover of putatively pathogenic, root‐associated fungi (hereafter, ‘pathogens’) in communities invaded by exotic plants, we conducted a two‐phase plant–soil feedback experiment: a monoculture experiment with native and exotic plants grown alone and a multi‐species, community‐level experiment that ranged in the extent of exotic dominance. We used next‐generation sequencing to characterise sharing of pathogens between native and exotic plants in communities. Exotic plants outperformed natives in communities, despite harbouring higher relative abundance of generalist pathogens. The higher generalism of pathogens supported by exotic plants made them more prone to be shared with natives. The proportion of pathogens shared between exotic and native plants in communities correlated with reduced competitive ability of native compared with exotic plants. Synthesis: These data suggest that exotic plants host more generalist pathogens that are shared with native plants, which may confer an indirect benefit to exotic over native plants through apparent competition.

Root traits and soil legacies drive species competition outcomes

Abstract Plant competition can be affected by plant functional traits but also by differences in fitness mediated by soil microbes. Climatic conditions such as drought further influence plant competition. Yet little is known about how soil microbes and drought interact with plant species that have distinct root traits and how this influences plant competition outcomes. We grew three plant species that co‐occur in temperate grasslands in China (Stipa krylovii, Artemisia frigida, Agropyron cristatum) in monocultures and mixtures and subjected the plant combinations to five soil inocula (root‐associated soil of S. krylovii, A. frigida, A. cristatum, an equal mixture of the three root zone soils and sterilized soil) as well as to a drought treatment. The relative change in plant biomass was used to determine plant competition outcomes. The three species exhibited clear differences in competitive abilities with A. cristatum > S. krylovii > A. frigida, and soil inocula or the drought treatment did not change the order. The relative yield (RY) of plants was affected by soil inocula, drought and plant arrangement. The strongest competitor, A. cristatum, with high total root length, root surface area and root volume experienced more negative biotic feedback, and drought enhanced the magnitude of these negative effects. On the contrary, the most inferior competitor, A. frigida, with high specific root length tended to have neutral or positive biotic feedback, and drought had no effect. Furthermore, the RY and fitness difference (reflected as the competitive ability in the mixture) of the three species were differentially influenced by root traits and plant–soil feedback. RY of A. cristatum could be predicted by the feedback effect in the mixture, and the fitness difference was mainly related to root traits. Both RY and fitness differences of A. frigida (the weakest competitor) could be predicted by root trait differences and feedback effects. Differences in root traits were the best predictors of the intermediate competitor S. krylovii. Our study shows that competition outcomes of co‐existing species depend on root traits and species‐specific PSF effects in mixture. Future work should examine the mechanisms that explain how plant competition and soil microbial heterogeneity act in conjunction with climate change in influencing plant coexistence. Read the free Plain Language Summary for this article on the Journal blog.

Soil Legacies of Tree Species Composition in Mature Forest Affect Tree Seedlings’ Performance

AbstractTrees affect the biotic and abiotic properties of the soil in which they grow. Tree species-specific effects can persist for a long time, even after the trees have been removed. We investigated to what extent such soil legacies of different tree species may impact tree seedlings in their emergence and growth. We performed a plant–soil feedback experiment, using soil that was conditioned in plots that vary in tree species composition in Białowieża Forest, Poland. Soil was taken from plots varying in proportion of birch, hornbeam, pine, and oak. In each soil, seeds of the same four target species were sown in pots. Seedling emergence and growth were monitored for one growing season. To further explore biotic implications of soil legacies, ectomycorrhizal root tip colonization of oak, a keystone forest species, was determined. We found no effect of soil legacies of tree species on the emergence measures. We, however, found a clear negative effect of pine legacies on the total biomass of all four seedling species. In addition, we found relationships between the presence of pine and soil fertility and between soil fertility and root tip colonization. Root tip colonization was positively correlated with the biomass of oak seedlings. We conclude that tree species can leave legacies that persist after that species has been removed. These legacies influence the growth of the next generation of trees likely via abiotic and biotic pathways. Thus, the choice of species in today’s forest may also matter for the structure and composition of future forests.

Seasonal drought treatments impact plant and microbial uptake of nitrogen in a mixed shrub grassland on the Colorado Plateau.

For many drylands, both long- and short-term drought conditions can accentuate landscape heterogeneity at both temporal (e.g., role of seasonal patterns) and spatial (e.g., patchy plant cover) scales. Furthermore, short-term drought conditions occurring over one season can exacerbate long-term, multidecadal droughts or aridification, by limiting soil water recharge, decreasing plant growth, and altering biogeochemical cycles. Here, we examine how experimentally altered seasonal precipitation regimes in a mixed shrub grassland on the Colorado Plateau impact soil moisture, vegetation, and carbon and nitrogen cycling. The experiment was conducted from 2015 to 2019, during a regional multidecadal drought event, and consisted of three precipitation treatments, which were implemented with removable drought shelters intercepting ~66% of incoming precipitation including: control (ambient precipitation conditions, no shelter), warm season drought (sheltered April-October), and cool season drought (sheltered November-March). To track changes in vegetation, we measured biomass of the dominant shrub, Ephedra viridis, and estimated perennial plant and ground cover in the spring and the fall. Soil moisture dynamics suggested that warm season experimental drought had longer and more consistent drought legacy effects (occurring two out of the four drought cycles) than either cool season drought or ambient conditions, even during the driest years. We also found that E. viridis biomass remained consistent across treatments, while bunchgrass cover declined by 25% by 2019 across all treatments, with the earliest declines noticeable in the warm season drought plots. Extractable dissolved inorganic nitrogen and microbial biomass nitrogen concentrations appeared sensitive to seasonal drought conditions, with dissolved inorganic nitrogen increasing and microbial biomass nitrogen decreasing with reduced soil volumetric water content. Carbon stocks were not sensitive to drought but were greater under E. viridis patches. Additionally, we found that under E. viridis, there was a negative relationship between dissolved inorganic nitrogen and microbial biomass nitrogen, suggesting that drought-induced increases in dissolved inorganic nitrogen may be due to declines in nitrogen uptake from microbes and plants alike. This work suggests that perennial grass plant-soil feedbacks are more vulnerable to both short-term (seasonal) and long-term (multiyear) drought events than shrubs, which can impact the future trajectory of dryland mixed shrub grassland ecosystems as drought frequency and intensity will likely continue to increase with ongoing climate change.

Plant-soil microbe feedbacks depend on distance and ploidy in a mixed cytotype population of Larrea tridentata.

Theory predicts that mixed ploidy populations should be short-lived due to strong fitness disadvantages for the rare ploidy. However, mixed ploidy populations are common, suggesting that the fitness costs for rare ploidies are counterbalanced by ecological benefits that emerge when rare. We investigated whether differences in ecological interactions with soil microbes help to maintain a tetraploid-hexaploid population of Larrea tridentata (creosote bush) in the Sonoran Desert, California, United States, where prior work documented ploidy-specific root-associated microbes. We used a plant-soil feedback (PSF) experiment to test whether host-specific soil microbes can alter the outcomes of intraploidy vs. interploidy competition. Host-specific soil microbes can build up over time; thus, distance from a host plant can affect the fitness of nearby plants. Seedlings grown in soils from near plants of a different ploidy produced greater biomass relative to seedlings grown in soils from near plants of the same ploidy. Moreover, seedlings grown in soils from near plants of a different ploidy produced more biomass than those grown in soils that were farther from plants of a different ploidy. These results suggest that the ecological consequences of PSF may facilitate the persistence of mixed ploidy populations. This is the first evidence, to our knowledge, that is consistent with plant-soil microbe feedback as a viable mechanism to maintain the coexistence of multiple ploidy levels in a single population.

Drought-induced assembly of rhizosphere mycobiomes shows beneficial effects on plant growth.

We have presented a framework to integrate the shifts in community assembly processes with plant-soil feedback during drought stress. We found that environmental filtering and host plant selection exert influence on the rhizospheric fungal community assembly, and the re-assembled community has great potential to alleviate plant drought stress. Our study proposes that future research should incorporate ecology with plant, microbiome, and molecular approaches to effectively harness the rhizospheric microbiome for enhancing the resilience of crop production to drought.

Restoration of degraded alpine meadows from the perspective of plant–soil feedbacks

Restoration of degraded alpine meadows from the perspective of plant–soil feedbacks

Vegetative growth drives the negative effects of an invasive species on resident community diversity and is not limited by plant-soil feedbacks: A temporal assessment.

Many pathways of invasion have been posited, but ecologists lack an experimental framework to identify which mechanisms are dominant in a given invasion scenario. Plant-soil feedbacks (PSFs) are one such mechanism that tend to initially facilitate, but over time attenuate, invasive species' impacts on plant diversity and ecosystem function. PSFs are typically measured under greenhouse conditions and are often assumed to have significant effects under field conditions that change over time. However, direct tests of PSFs effects in natural settings and their change over time are rare. Here we compare the role of PSFs with the effects of biomass in limiting the dominance of an invasive species and impacts on resident species diversity. We characterized the effects of the invader Bromus inermis (Leyss.) on native plant communities over time and measured changes in its conspecific PSFs and vegetative growth to understand their integrated effects on community diversity. To do so, we combined data from a 6-year field study documenting the rate and impacts of invasion with a short-term greenhouse experiment quantifying PSF as a function of time since invasion in the field. We found that the nature and strength of B. inermis PSFs did not change over time and were not mediated by soil microbial communities. Though PSFs impacted B. inermis reproduction, they did not sufficiently limit vegetative growth to diminish the negative impacts of B. inermis biomass on native species. B. inermis experienced the full strength of its negative PSFs immediately upon invasion, but they were ineffective at reducing B. inermis vigor to facilitate the recovery of the native plant community. We recommend that conservation efforts focus on limiting B. inermis vegetative growth to facilitate community recovery.

Intraspecific variation in Janzen-Connell effect is mediated by stress and plant-soil feedbacks.

Janzen-Connell (JC) effects, hypothesized to be partially driven by negative plant-soil feedbacks (PSFs), are considered to be a key mechanism that regulates tropical forest plant diversity and coexistence. However, intraspecific variation in JC effects may weaken this mechanism, with the strength of PSFs being a potentially key variable process. We conducted a manipulated experiment with seedlings from two populations of Pometia pinnata (Sapindaceae), a tropical tree species in southwest China. We aimed to measure the intraspecific difference in PSF magnitude caused by inoculating the soil from different P. pinnata source populations and growing seedlings under differing light intensity and water availability treatments, and at varying plant densities. We found negative PSFs for both populations with the inoculum soil originating from the same sites, but PSFs differed significantly with the inoculum soil from different sites. PSF strength responded differently to biotic and abiotic drivers; PSF strength was weaker in low moisture and high light treatments than in high moisture and low light treatments. Our study documents intraspecific variation in JC effects: specifically, P. pinnata have less defenses to their natively-sourced soil, but are more defensive to the soil feedbacks from soil sourced from other populations. Our results imply that drought and light intensity tended to weaken JC effects, which may result in loss of species diversity with climate change.

Assessing the Influence of Soil Composition on Plant Growth and Development in USA

Purpose: The aim of the study was to assess the influence of soil composition on plant growth and development. Methodology: This study adopted a desk methodology. A desk study research design is commonly known as secondary data collection. This is basically collecting data from existing resources preferably because of its low cost advantage as compared to a field research. Our current study looked into already published studies and reports as the data was easily accessed through online journals and libraries. Findings: Assessing soil composition's influence on plant growth and development reveals crucial insights. Soil pH affects nutrient availability, with acidic soils limiting essential nutrient uptake. Organic matter content enhances soil structure and microbial activity, promoting nutrient cycling. Soil texture influences water infiltration and root penetration, impacting plant establishment. Toxic elements like heavy metals inhibit plant growth by disrupting physiological processes. Unique Contribution to Theory, Practice and Policy: Liebig's law of the minimum, optimal foraging theory & soil-plant feedback theory may be used to anchor future studies on the soil composition on plant growth and development. Develop soil management guidelines tailored to specific agro ecological contexts, taking into account soil properties, climate conditions, and crop requirements to optimize nutrient availability and uptake. Integrate soil health indicators and ecosystem services assessments into agricultural policies and land use planning frameworks to promote sustainable soil management practices and mitigate environmental degradation.

Effects of plant-soil feedbacks on the invasion and competitiveness of Aegilops tauschii

The interaction between plants and soil is an important aspect that affects the invasive ability of foreign plants and the invasiveness of ecosystems. The study on plant soil feedback of A. tauschii can provide reference for its invasion mechanism. Firstly, the effects of A. tauschii on the nutrients and enzyme activities of invaded soil were investigated; Secondly, in conjunction with soil sterilization, pot experiments were conducted using the De Wit substitution method to investigate the impact of different degree invasive soils on the development of A. tauschii and its interaction with wheat. The results showed that the invasion of A. tauschii significantly increased soil organic matter, soluble phosphorus and soluble potassium, while also causing a significant decrease in the concentration of nitrate nitrogen. And according to the changes of morphological and biomass indicators of A. tauschii, the results of two-way ANOVAs showed that the invaded soil and its microbiota have a positive feedback effect on the growth of A. tauschii. Finally, it can be seen from the value of the competition balance index, the competition ability of A. tauschii in different invasion degree soil is greater than that of wheat whether the soil invaded by A. tauschii had undergone sterilization treatment or not. In conclusion, the invasion potential of A. tauschii is not only derived from its strong competitiveness, but also may be related to the soil conditions.

Self‐help or self‐sabotage? A common invader's soil legacy does not impede, and may facilitate, potential competitors

AbstractIntroduced plants create lasting abiotic and biotic changes to soil that are increasingly implicated as major drivers of invasion success, yet models for investigating patterns in the impacts of these soil legacies are lacking. Here, we used Bohemian knotweed (Reynoutria × bohemica) to present a novel approach for testing whether accounting for diverse evolutionary drivers can clarify how an invader's soil legacy mediates its success. For most tested species, R. × bohemica's soil legacy did not influence seedling performance irrespective of its evolutionary history or the evolutionary novelty of species that share its soil. However, we found the relative importance of legacy effects versus other mechanisms of invader dominance depends on the life stage of the plants growing in soil conditioned by the invader. While the effects of R. × bohemica's soil legacy on seedling performance were context dependent, emergence was, on average, higher in R. × bohemica‐conditioned soils than in unconditioned soils. Thus, contrary to theory, R. × bohemica's soil legacy not only fails to impede potential competitors but may also facilitate them. Our results are particularly powerful: we detected novel insights in the face of biological complexity, even for R. × bohemica, which likely does not depend on heterospecific plant–soil feedbacks for its competitive dominance in its introduced range. Designing rigorous investigations that explore how plant species' evolutionary histories mediate an invader's soil legacy across diverse abiotic environments, as shown here, is key for producing insights in other systems.

Effects of riparian pioneer plants on soil aggregate stability: Roles of root traits and rhizosphere microorganisms

Pioneer plants are vital in stabilizing soil structure while restoring reservoir drawdown areas. However, uncertainties persist regarding the mechanism of pioneer plants to soil stability in these delicate ecosystems. This study aims to unravel the plant-soil feedback mechanisms from the roles of root traits and rhizosphere microorganisms. We conducted a mesocosm experiment focusing on four common pioneer plants from the drawdown area of Three Gorges Reservoir, China. Using the wet sieving methodology, trait-based approach and high-throughput sequencing technology, we explored soil aggregate stability parameters, plant root traits and rhizosphere microbial communities in experimental plant groups. The interacting effect of pioneer plant species richness, root traits, and rhizosphere microbial communities on soil aggregate stability was quantified by statistical and machine-learning models. Our results demonstrate that diverse pioneer plant communities significantly enhance soil aggregate stability. Notably, specific species, such as Cynodon dactylon (L.) Pers. and Xanthium strumarium L., exert a remarkably strong influence on soil stability due to their distinctive root traits. Root length density (RLD) and root specific surface area (RSA) were identified as crucial root traits mediating the impact of plant diversity on soil aggregate stability. Additionally, our study highlights the link between increased rhizosphere fungal richness, accompanied by plant species richness, and enhanced soil aggregate stability, likely attributable to elevated RLD and RSA. These insights deepen our understanding of the role of pioneer vegetation in soil structure and stability, providing valuable implications for ecological restoration and management practices in reservoir drawdown areas.

Plant functional traits modulate the effects of soil acidification on above- and belowground biomass

Abstract. Atmospheric sulfur (S) deposition has been increasingly recognized as a major driver of soil acidification. However, little is known about how soil acidification influences above- and belowground biomass by altering leaf and root traits. We conducted a 3-year S-addition experiment to simulate soil acidification in a meadow. Grass (Leymus chinensis (Trin.) Tzvelev) and sedge (Carex duriuscula C.A.Mey) species were chosen to evaluate the linkage between plant traits and biomass. Sulfur addition led to soil acidification and nutrient imbalances. Soil acidification decreased specific leaf area (SLA) but increased leaf dry-matter content (LDMC) in L. chinensis, showing a conservative strategy and thus suppressing aboveground instead of belowground biomass. However, in C. duriuscula, soil acidification increased plant height and root nutrients (N, P, S, and Mn), favoring competition for natural resources through enhanced above- and belowground biomass, i.e., adoption of an acquisitive strategy. Increased soil acidity resulted in an overall reduction in aboveground community biomass by 3 %–33 %, but it led to an increase in community root biomass by 11 %–22 % due to upregulation as a result of higher soil nutrient availability. Our results demonstrate that both above- and belowground plant biomass is affected by S-induced acidification. Understanding the linkage between plant biomass and functional traits contributes to a better understanding of plant–soil feedback in grassland ecosystems.

Biotic plant-soil feedbacks alone do not explain why more diverse plant communities are less invasible

Biotic plant-soil feedbacks alone do not explain why more diverse plant communities are less invasible

Intraspecific plant-soil feedback in four tropical tree species is inconsistent in a field experiment.

Soil microbes can influence patterns of diversity in plant communities via plant-soil feedbacks. Intraspecific plant-soil feedbacks occur when plant genotype leads to variations in soil microbial composition, resulting in differences in the performance of seedlings growing near their maternal plants versus seedlings growing near nonmaternal conspecific plants. How consistently such intraspecific plant-soil feedbacks occur in natural plant communities is unclear, especially in variable field conditions. In an in situ experiment with four native tree species on Barro Colorado Island (BCI), Panama, seedlings of each species were transplanted beneath their maternal tree or another conspecific tree in the BCI forest. Mortality and growth were assessed at the end of the wet season (~4 months post-transplant) and at the end of the experiment (~7 months post-transplant). Differences in seedling performance among field treatments were inconsistent among species and eroded over time. Effects of field environment were detected at the end of the wet season in two of the four species: Virola surinamensis seedlings had higher survival beneath their maternal tree than other conspecific trees, while seedling survival of Ormosia macrocalyx was higher under other conspecific trees. However, these differences were gone by the end of the experiment. Our results suggest that intraspecific plant-soil feedbacks may not be consistent in the field for tropical tree species and may have a limited role in determining seedling performance in tropical tree communities. Future studies are needed to elucidate the environmental and genetic factors that determine the incidence and direction of intraspecific plant-soil feedbacks in plant communities.

Effect of plant-soil system on the restoration of community stability after wildfire in the northeast margin of Qinghai-Tibet plateau

Wildfires, as an environmental filter, are pivotal ecological disturbances that reshape plant communities and soil dynamics, playing a crucial role in regulating biogeographic patterns and ecosystem services. In this study, we aim to explore the effects of wildfires on forest ecosystems, specifically focusing on the plant-soil feedback mechanisms within the northeastern margin of the Qinghai-Tibet Plateau (QTP). Utilizing Partial Least Squares Path Modeling (PLS-PM), we investigated the interrelationships among soil physicochemical properties, enzyme activities, species diversity, and community stability at varying post-fire recovery stages (5, 15, and 23 years). Results indicated that in the early recovery stages, rapid changes in soil properties such as decreased pH (p < 0.001) and increased nutrient availability facilitate the emergence of early successional species with high resource utilization traits. As the ecosystem evolved toward a climax community, the soil and vegetation exhibit increased stability. Furthermore, soil enzyme activities displayed dynamic patterns that corresponded with changes in soil nutrient content, directly influencing the regeneration and diversity of plant communities. Importantly, our study documented a transition in the influence of soil properties on community stability from direct positive effects in initial recovery phases to negative impacts in later stages, while indirect benefits accrue through increased species diversity and enzyme activity. Vegetation composition and structure changed dynamically with recovery time during community succession. Plant nutrient absorption and accumulation affected nutrient dynamics in the soil, influencing plant regeneration, distribution, and diversity. Our results underscore the complex interactions between soil and vegetation that drive the recovery dynamics post-wildfire, highlighting the resilience of forest ecosystems to fire disturbances. This study contributes to the understanding of post-fire recovery processes and offers valuable insights for the management and restoration of fire-affected forest ecosystems.

Why are graminoid species more dominant? Trait-mediated plant-soil feedbacks shape community composition.

Species traits may determine plant interactions along with soil microbiome, further shaping plant-soil feedbacks (PSFs). However, how plant traits modulate PSFs and, consequently, the dominance of plant functional groups remains unclear. We used a combination of field surveys and a two-phase PSF experiment to investigate whether forbs and graminoids differed in PSFs and in their trait-PSF associations. When grown in forb-conditioned soils, forbs experienced stronger negative feedbacks, while graminoids experienced positive feedbacks. Graminoid-conditioned soil resulted in neutral PSFs for both functional types. Forbs with thin roots and small seeds showed more-negative PSFs than those with thick roots and large seeds. Conversely, graminoids with acquisitive root and leaf traits (i.e., thin roots and thin leaves) demonstrated greater positive PSFs than graminoids with thick roots and tough leaves. By distinguishing overall and soil biota-mediated PSFs, we found that the associations between plant traits and PSFs within both functional groups were mainly mediated by soil biota. A simulation model demonstrated that such differences in PSFs could lead to a dominance of graminoids over forbs in natural plant communities, which might explain why graminoids dominate in grasslands. Our study provides new insights into the differentiation and adaptation of plant life-history strategies under selection pressures imposed by soil biota.

Negative conspecific plant-soil feedback on alien plants co-growing with natives is partly mitigated by another alien

Abstract Background and aims Naturalized alien and native plants can impact each other directly when they grow next to each other, but also indirectly through their soil legacies. These alien-native interactions can also be modified by the presence of a third alien or native species. However, it is unknown how the performance of co-growing alien and native species is affected by their soil legacies and by the presence of an additional species. Methods In our two-phase plant-soil-feedback experiment, soils were first conditioned by eight herbaceous species, four of which are naturalized and four of which are native to Germany. We then grew all 16 pairwise alien-native species combinations on soil conditioned by the respective alien or native species, on a mixture of soils conditioned by both species or on control soil. Each pair of test plants was grown on these soils without or with an additional alien or native species. Results Soil conditioning, and particularly conspecific soil conditioning, reduced growth of the alien and native test plants. The addition of another species also reduced growth of the test plants. However, the negative conspecific soil-legacy effect on alien test plants was reduced when the additional species was also alien. Conclusion The negative conspecific plant-soil feedback for alien and native plants in our study could promote their coexistence. However, as partial alleviation of negative conspecific effects on alien plants occurred with an additional alien species, the chances of coexistence of alien and native species might decrease when there are multiple alien species present.

Reduced belowground allocation of freshly assimilated C contributes to negative plant-soil feedback in successive winter wheat rotations

Abstract Aims Successive winter wheat (WW) rotations are associated with yield reduction, often attributed to the unfavorable soil microbes that persist in the soil through plant residues. How rotational positions of WW affect the allocation of freshly assimilated carbon (C), an energy source for soil microbes, above and belowground remains largely unknown. Methods A 13CO2 pulse labeling rhizotron experiment was conducted in the greenhouse to study freshly fixed C allocation patterns. WW was grown in soil after oilseed rape (W1), after one season of WW (W2), and after three successive seasons of WW (W4). We used an automatic manifold system to measure excess 13C of soil respiration at six depths and five different dates. Excess 13C was also measured in dissolved organic C (DOC), microbial and plant biomass pools. Results There was a strong yield decline in successive WW rotations accompanied by distinct changes in root growth. Higher excess 13C of soil respiration was measured in W1 compared to W4, especially in the topsoil during at later growth stages. Higher excess 13C of the DOC and the microbial biomass was also traced in W1 and W4 compared to W2. Less 13C was taken up by successive WW rotations. Conclusions Our study demonstrates a mechanism through which the rotational position of WW affects the allocation of freshly assimilated C above and belowground. WW after oilseed rape sustains belowground allocation of freshly assimilated C for a longer time than successively grown WW and incorporates more of this C to its biomass.