Belowground Biomass Allocation Research Articles

Winter Grazing, Not Fencing or Unicast, Promotes Stability of Microbial Community and Function in the Qilian Mountains of Qinghai‐Xizang Plateau

ABSTRACTIn alpine meadows, microorganisms are essential to sustain the stability of terrestrial geochemical processes and vegetation–soil–microbial systems. The present study in order investigate how various management measures impact the microbial communities' composition and functionality, we utilize metagenomic sequencing techniques to examinate the composition and function of soil microbial communities in the southern Qilian Mountains of the Qinghai‐Xizang Plateau in response to the management practices of fencing enclose (FE), winter grazing (WG), transition zone between natural and artificial grasslands (TZ), and artificial unicast oats (AU). Vegetation diversity and soil physicochemical characteristics were dramatically altered by the management measures. The prokaryotic community structure was considerably similar in FE and WG, as well as in TZ and AU. Near‐natural (FE) and artificial establishment (AU) disturbances changed the fungal community structure. Enzymes related to carbon metabolism did not respond significantly to the management measures, whereas those related to nitrogen metabolism did not respond significantly in TZ and AU. The relative abundance of enzymes participating in nitrogen metabolism was higher under TZ and AU than under FE and WG. We concluded that grassland management measures altered the structure of aboveground graminoid and leguminous vegetation communities and belowground biomass allocation, resulting in changes in K uptake, causing striking changes in the structure of fungal communities and nitrogen‐metabolizing enzymes; moderate disturbance (WG) was beneficial for maintaining the stability of microbial communities in alpine grasslands.

Trait-Mediated Variation in Seedling Performance in Costa Rican Successional Forests: Comparing Above-Ground, Below-Ground, and Allocation-Based Traits.

The interspecific relationship between functional traits and tree seedling performance can be inconsistent, potentially due to site-to-site or microsite variation in environmental conditions. Studies of seedling traits and performance often focus on above-ground traits, despite the importance of below-ground resource acquisition and biomass allocation to above versus below-ground functions. Here we investigate how varying environmental conditions across sites induce intraspecific variation in organ-level (above-ground, below-ground) and biomass allocation traits, affecting interspecific relationships between these traits and seedling performance. We analyzed trait expression for 12 organ-level and three allocation traits and their relationships with height growth (1716 seedlings) and mortality (15,862 seedlings) for 26 tree species across three sites along a forest successional gradient in Costa Rica. We found significant intraspecific differences across sites in all allocation traits, but only in three of seven above-ground and three of five below-ground organ-level traits. Allocation traits were better predictors of seedling performance than organ-level traits. Relationships between allocation traits and both growth and mortality varied among all sites, but for organ-level traits, only relationships with growth varied among sites. These results underscore that biomass allocation plays a key role in the earliest life stages of trees and that site-specific conditions can influence how functional traits mediate seedling establishment during succession.

The above and the belowground nitrogen allocation strategy of Scirpus mariqueter based on 15N isotope tracing along an elevation gradient and its significance for coastal wetlands restoration

AbstractBackgroundEcological restoration of coastal wetlands has become particularly urgent worldwide as wetland areas have declined dramatically over the past two decades.AimsTo understand the nitrogen allocation strategy of Scirpus mariqueter (S. mariqueter) and provide theory support for future wetland management and restoration.MethodsThe study investigated the response mechanism of S. mariqueter to altitude spatial changes in Nanhui Dongtan from 2017 to 2019 using remote sensing imagery and field surveys. The ecological adaptability of S. mariqueter at different elevations (denoted as A, B, and C for elevations 2.40, 3.15, and 3.49 m, respectively) was analyzed through 15N stable isotope tracing technology. In July and September 2020, 15N‐enriched urea solution was uniformly sprayed onto the leaf surfaces of S. mariqueter at different sites. Plant samples were collected at the end of July and September, and the aboveground, belowground, seed, and rhizome biomass were measured, followed by 15N isotope tests.Results(1) From 2017 to 2019, the biomass of S. mariqueter significantly increased in the elevation change of 0.44–0.48 m, with the maximum aboveground biomass increase of 488.70 g dm−2. The density also increased significantly in the elevation change of 0.13–0.43 m, peaking at 674.02 plants m−2; (2) during the growing period, the biomass of A, B, and C increased. The aboveground portion of the 15N allocation rate accounted for 74%–84%. The belowground portion of the 15N allocation rate positively correlated with elevation; (3) during the reproductive period, elevation positively correlated with the 15N distribution rate of seeds and corms, as well as the biomass allocation rate of seeds and aboveground portions. The 15N allocation rate of the corms was higher than that of seeds. Additionally, elevation exhibited a negative correlation with belowground biomass allocation rate. (4) Point A has the highest difference of above and belowground biomass proportion, and 15N isotope allocation. Area of point A is the critical area affecting vegetation expansion and should be paid more attention in the future work of coastal management and restoration.ConclusionThere is an adaption strategy of S. mariqueter that affects the plant‐soil interaction and the biogeomorphological development process along the elevation gradient.

Biomass Allocation of China’s Forests as Indicated by a Literature-Based Allometry Database

Allometry reflects the quantitative relationship between the allocation of resources among different organs. Understanding patterns of forest biomass allocation is critical to comprehending global climate change and the response of terrestrial vegetation to climate change. By collecting and reorganizing the existing allometric models of tree species in China, we established a database containing over 3000 empirical allometric models. Based on this database, we analyzed the model parameters and the effect of climate on forest biomass allocation under the context of ‘optimal allocation theory’. We showed that (1) the average and median exponent of power functions for above-ground biomass were 2.344 and 2.385, respectively, which significantly deviated from the theoretical prediction of 2.667 by metabolic theory (p < 0.01). (2) The parameters of the allometric model were not constant, and not significantly correlated with temperature, precipitation, latitude, and elevation (p > 0.05), but were more closely related to individual size (p < 0.01). (3) Among different types of forests, the proportion of above-ground biomass in tropical rainforests and subtropical evergreen rainforests was significantly higher than that in temperate forests and boreal forests (p < 0.05). The proportion of trunk and branch biomass allocated to tropical rainforest was significantly higher than that of boreal forest (p < 0.05), while the proportion of root and leaf biomass allocated to tropical rainforest was significantly lower than that of boreal forest (p < 0.05). (4) The abiotic environment plays a crucial role in determining the allocation of plant biomass. The ratio of below-ground/above-ground biomass is significantly and negatively correlated with both temperature and rainfall (p < 0.01), and significantly and positively correlated with altitude and latitude (p < 0.01). This means that as temperature and rainfall increase, there is a decrease in the amount of biomass allocated to below-ground structures such as roots. On the other hand, as altitude and latitude increase, there is an increase in below-ground biomass allocation. These findings highlight the importance of considering the influence of abiotic factors on plant growth and development.

Climate Factors Influence Above- and Belowground Biomass Allocations in Alpine Meadows and Desert Steppes through Alterations in Soil Nutrient Availability.

Biomass is a direct reflection of community productivity, and the allocation of aboveground and belowground biomass is a survival strategy formed by the long-term adaptation of plants to environmental changes. However, under global changes, the patterns of aboveground-belowground biomass allocations and their controlling factors in different types of grasslands are still unclear. Based on the biomass data of 182 grasslands, including 17 alpine meadows (AMs) and 21 desert steppes (DSs), this study investigates the spatial distribution of the belowground biomass allocation proportion (BGBP) in different types of grasslands and their main controlling factors. The research results show that the BGBP of AMs is significantly higher than that of DSs (p < 0.05). The BGBP of AMs significantly decreases with increasing mean annual temperature (MAT) and mean annual precipitation (MAP) (p < 0.05), while it significantly increases with increasing soil nitrogen content (N), soil phosphorus content (P), and soil pH (p < 0.05). The BGBP of DSs significantly decreases with increasing MAP (p < 0.05), while it significantly increases with increasing soil phosphorus content (P) and soil pH (p < 0.05). The random forest model indicates that soil pH is the most important factor affecting the BGBP of both AMs and DSs. Climate-related factors were identified as key drivers shaping the spatial distribution patterns of BGBP by exerting an influence on soil nutrient availability. Climate and soil factors exert influences not only on grassland biomass allocation directly, but also indirectly by impacting the availability of soil nutrients.

Climate Factors Affect Above–Belowground Biomass Allocation in Broad-Leaved and Coniferous Forests by Regulating Soil Nutrients

The allocation of plant biomass above and below ground reflects their strategic resource utilization, crucial for understanding terrestrial carbon flux dynamics. In our comprehensive study, we analyzed biomass distribution patterns in 580 broadleaved and 345 coniferous forests across China from 2005 to 2020, aiming to discern spatial patterns and key drivers of belowground biomass proportion (BGBP) in these ecosystems. Our research revealed a consistent trend: BGBP decreases from northwest to southeast in both forest types. Importantly, coniferous forests exhibited significantly higher BGBP compared to broadleaved forests (p &lt; 0.001). While precipitation and soil nutrients primarily influenced biomass allocation in broadleaved forests, temperature and soil composition played a pivotal role in coniferous forests. Surprisingly, leaf traits had a negligible impact on BGBP (p &gt; 0.05). Climatic factors, such as temperature and rainfall, influenced biomass partitioning in both strata by altering soil nutrients, particularly soil pH. These findings provide valuable insights into understanding carbon sequestration dynamics in forest ecosystems and improving predictions of the future trajectory of this critical carbon cycle component.

Climate and breeding determined below-ground biomass allocation strategy in wheat

Farmland covers about 12% of the ice-free terrestrial surface, and crop below-ground biomass allocation plays a pivotal role in the sequestration of atmospheric carbon. Nevertheless, our understanding of the integrated response of crop belowground biomass allocation strategy to concomitant variation in key environmental factors related to water and nutrient availability remains limited. Furthermore, it is imperative to conduct further analysis to elucidate the influence of breeding efforts geared toward improving crop yield on the below-ground biomass allocation strategy. Here we compiled a global wheat root mass fraction (RMF) database with 221 observations from 39 studies to analyze the integrated response of RMF to climate, soil properties and filed managements, and the relationship of RMF and yield. We identified an aridity index (AI) threshold of 0.44 for wheat below-ground biomass allocation strategy, based on the effect of long-term climate on plow-layer soil organic carbon and field water capacity (FWC). The negative effect of nitrogen fertilizer rate on RMF along the AI gradient did not significantly change but the main mechanism shifted from indirect to direct effects. For AI < 0.44, the response of RMF to change of water availability was relatively conservative with the insignificant effects of FWC on RMF. However, RMF was highly sensitive to precipitation distribution during the growth stage because RMF decreased as the ratio of precipitation during the reproductive stage to the growth stage increased. We did not observe a significant correlation between RMF and yield within the main range of RMF so a large amount of intraspecific variation in belowground biomass allocation strategies had not been filtered out by breeding. For AI ≥ 0.44, RMF was highly sensitive to water resource availability as RMF significantly decreased with an increase in FWC. Yield decreased significantly with increasing RMF within the main range of RMF, resulting in highly consistent intraspecific belowground biomass allocation strategies under the pressure of breeding. The results suggested long-term climate condition and breeding played critical filters in wheat below-ground biomass allocation strategy and provided a more accurate prediction of RMF for global carbon cycle models and crop growth models.

Link between the aboveground and belowground biomass allocation with growing of Tamarix sp. seedlings in the hinterland of Taklimakan Desert, China.

The morphological characteristics and biomass allocation can reflect plant adaptive strategies to the environment. Tamarix sp. is an excellent shrub species used for windbreaks and fixing sand in the desert of northwest China. The successful establishment of Tamarix sp. seedlings and their growth into mature individuals require their adaptation to various environmental conditions, which is the key to naturally regenerating the Tamarix population. To clarify the root morphological characteristics, leaf structural characters, and biomass allocation of Tamarix sp. seedlings in response to drought conditions, we took the Tamarix sp. seedlings at the Daryaboyi oasis in the hinterland of Taklimakan Desert as the object of study, analyzed rooting depth, root dry weight (RDW), specific root length (SRL), root surface area (RA), specific root area (SRA), leaf area (LA), specific leaf area (SLA) and root: shoot ratio (R:S ratio). The gravimetric soil water content varied from 5.80% to 25.84% in this study area. The taproots of Tamarix sp seedlings with small basal stem diameters were shallower and had few lateral root branches and Tamarix sp. seedlings with large basal stem diameters had more obvious taproots and lateral roots. With the growth of Tamarix sp. seedlings, the taproot deepened, and the values ranged from 4.5 cm to 108.0 cm; the SRL, SRA, and SLA decreased, and the ranges of the values were 28.92-478.79 cm·g-1, 1.07-458.50 cm2·g-1, and 24.48-50.7 cm2·g-1; the RDW, RA, and LA increased, the ranges of the values were 0.16-21.34 g, 3.42-328.04 cm2, and 2.41-694.45 cm2; the more biomass was allocated to the aboveground parts, and the mean R: S ratio was 0.76. In better soil water conditions, the root growth rate decreased as Tamarix sp. seedlings grew, and more biomass was allocated to the aboveground. This further showed that stable surface water is highly significant to the biomass allocation strategy of Tamarix sp. seedlings.

Plants alter their aboveground and belowground biomass allocation and affect community-level resistance in response to snow cover change in Central Asia, Northwest China

It is important to elucidate the changing distribution pattern of net primary productivity (NPP) to mechanistically understand the changes in aboveground and belowground ecosystem functions. In water-scarce desert environments, snow provides a crucial supply of water for plant development and the spread of herbaceous species. Yet uncertainty persists regarding how herbaceous plants' NPP allocation responds to variation in snow cover. The goal of this study was to investigate how variation in snow cover in a temperate desert influenced the NPP allocation dynamics of herbaceous species and their resistance to environmental change in terms aboveground and belowground productivity. In the Gurbantunggut Desert, wintertime snow cover depth was adjusted in plots by applying four treatments: snow removal (−S), ambient snow, double snow (+S), and triple snow (+2S). We examined their species richness, aboveground NPP (ANPP), belowground NPP (BNPP), and the resistance of ANPP and BNPP. We found that species diversity of the aboveground community increased significantly with increasing snow cover and decreased significantly Pielou evenness in plots. This resulted in greater ANPP with increasing snow cover; meanwhile, BNPP first increased and then decreased with increasing snow cover. However, this productivity in different soil layers responded differently to changed snow cover. In the 0–10 cm soil layer, productivity first rose and then declined, while it declined linearly in both the 10–20 cm and 20–30 cm soil layers, whereas in the 30–40 cm soil layer it showed an increasing trend. Belowground resistance would increase given that greater snow cover improved the BNPP in deeper soil and maintained the resource provisioning for plant growth, thus improving overall belowground stability. These results can serve as a promising research foundation for future work on how the functioning of desert ecosystems becomes altered due to changes in plant community expansion and, in particular, changes in snow cover driven by global climate change.

Vertical distribution patterns of community biomass, carbon and nitrogen content in grasslands on the eastern Qinghai–Tibet Plateau

The accumulation, transport and distribution of photosynthates within plants not only affect the ratio of shoot-to-root biomass, but also change the distribution pattern of nutrient elements in different organs and tissues. A full understanding of the allocation patterns of biomass and nutrient elements within plants, is helpful to strengthen our understanding of the above- and belowground ecosystem functions. However, at present, we do not know enough about the aboveground–belowground allocation of biomass, carbon (C) and nitrogen (N) contents, and, in particular, the corresponding vertical distribution pattern belowground and its environmental impact. At the community scale, the changes in vegetation community attributes measured in the field can reflect the long-term adaptation and subsequent adaptation strategies of vegetation to the regional environment and environmental changes. In this study, the vertical distribution patterns of community biomass and the carbon and nitrogen contents with soil depth in an alpine grassland on the eastern Qinghai–Tibet Plateau were investigated based on the transect method. Our results show the following: (1) Alpine grasslands can respond to differences in hydrothermal resource availability by changing biomass allocation patterns. In low-temperature and high-humidity environments, aboveground and belowground biomass allocation supports the isometric growth hypothesis. However, in environments with less soil moisture or high temperature and humidity, the aboveground and belowground biomass distributions support the allometric allocation hypothesis. (2) The vertical distribution patterns of the C content, N content and carbon: nitrogen ratio (C:N) within plants are different. From the leaves to the root system, with the increase in root depth, the C content demonstrated a vertical distribution pattern of first decreasing and then increasing, and the N% demonstrated a gradual decreasing distribution pattern, while the C:N ratio demonstrated an exponential increasing distribution pattern. (3) The change rate in C%, N% and C:N related to the soil depth was affected by environmental hydrothermal conditions. C% and C:N ratio changed more rapidly with root depth under low temperature and high humidity. The change rate of N% was mainly affected by temperature and changed more rapidly with root depth at low temperatures. (4) When exploring the relationship between the content of elements within plants, it is necessary to combine both the above- and belowground compartments. There was a nonlinear quadratic relationship between the carbon and nitrogen contents within plants.

Precipitation Dominates the Allocation Strategy of Above- and Belowground Biomass in Plants on Macro Scales.

The allocation of biomass reflects a plant's resource utilization strategy and is significantly influenced by climatic factors. However, it remains unclear how climate factors affect the aboveground and belowground biomass allocation patterns on macro scales. To address this, a study was conducted using aboveground and belowground biomass data for 486 species across 294 sites in China, investigating the effects of climate change on biomass allocation patterns. The results show that the proportion of belowground biomass in the total biomass (BGBP) or root-to-shoot ratio (R/S) in the northwest region of China is significantly higher than that in the southeast region. Significant differences (p < 0.05) were found in BGBP or R/S among different types of plants (trees, shrubs, and herbs plants), with values for herb plants being significantly higher than shrubs and tree species. On macro scales, precipitation and soil nutrient factors (i.e., soil nitrogen and phosphorus content) are positively correlated with BGBP or R/S, while temperature and functional traits are negatively correlated. Climate factors contribute more to driving plant biomass allocation strategies than soil and functional trait factors. Climate factors determine BGBP by changing other functional traits of plants. However, climate factors influence R/S mainly by affecting the availability of soil nutrients. The results quantify the productivity and carbon sequestration capacity of terrestrial ecosystems and provide important theoretical guidance for the management of forests, shrubs, and herbaceous plants.

Long-term nitrogen addition and drought altered root trait and plant biomass allocation of five herbs in early spring in a temperate forest

ABSTRACT In early spring, because the tree and shrub layers are still dormant, the absorption of nutrients by herbs is the key factor to maintain nutrients in temperate forest ecosystems, but it is not clear whether climate change will change the nutrient uptake strategy of herbs. Therefore, we measured the biomass, fine root morphology, and stoichiometry of five dominant herbs in response to environmental changes in early spring in a temperate forest in northeastern China. The results showed that (1) Fine root morphology had strong plasticity. Under nitrogen addition treatment, the herbs tended to change the fine root length to absorb soil nutrients and water. (2) Under drought treatment, the herbs distributed more biomass to the belowground. Maintaining a higher root-shoot ratio might increase the relative supply of water to the herbs. (3) Nitrogen addition significantly decreased the belowground biomass allocation, but increased the ratio of belowground nitrogen to phosphorus (N:P) and the root metabolic activity. (4) Ephemeral herbaceous fine root morphology was more sensitive to water and nitrogen than other herbaceous plants. Collectively, in the context of climate change, early spring herbs may respond to environmental pressure through self-regulation and change growth strategies in temperate forest ecosystems.

A response of biomass and nutrient allocation to the combined effects of soil nutrient, arbuscular mycorrhizal, and root-knot nematode in cherry tomato

IntroductionThe biomass and nutrient allocation strategies in plants are fundamental for predicting carbon storage and mineral and nutrient cycles in terrestrial ecosystems. However, our knowledge regarding the effects of multiple environmental factors on biomass and nutrient allocation remains limited.MethodsHere we manipulated soil composition (three levels), arbuscular mycorrhizal fungi inoculation (AMF, five levels), and root-knot nematode inoculation (RKN, two levels) using random block design to reveal the effects of these factors on biomass and nutrient allocation strategies of cherry tomato.Results and DiscussionOur results showed that biomass and nutrient allocation were affected by soil composition, AMF and RKN individually or interactively. The biomass and nutrient allocation in cherry tomato shows different adaptation strategies responded to the joint action of three factors. The reduction of soil nutrients increased belowground biomass allocation, and aboveground nitrogen and phosphorus concentration. AMF colonization increased aboveground biomass allocation and reproductive investment and promoted aboveground nitrogen and phosphorus inputs. Cherry tomato can mitigate the stress of RKN infection by investing more biomass and nutrients into belowground organs. Our study showed that plants can adjust their survival strategies by changing biomass and nutrient allocation to adapt to variation in soil abiotic and biotic factors. These findings contribute to our understanding of the adaptive processes of plant biomass and nutrient allocation strategies under multiple environmental factors.

Plant biomass allocation is mediated by precipitation use efficiency in arid and semiarid ecosystems

AbstractThe arid and semiarid grassland ecosystems are characterized by limited water resources and are generally vulnerable to climate change. Understanding how plants in arid and semiarid ecosystems respond to global climatic variation is crucial for ecological restoration under a changing climate. Although the effects of climate on aboveground biomass (AGB) and belowground biomass (BGB) have been widely explored, how AGB and BGB respond to climatic variation is seldom disentangled. There is still a need to understand how plant communities respond to global climate change. In this study, we conducted a transect survey across grasslands in Inner Mongolia to capture changes in AGB and BGB in plant communities. Then, we used structural equation modelling (SEM) to explore the relationship between environmental factors and the root:shoot ratio to understand how plant communities respond to ecological drought under global climate change. Our results showed that low precipitation use efficiency (LPUE) results in a high root:shoot (HRS) ratio, and BGB was more sensitive to environmental changes. By contrast, high precipitation use efficiency (HPUE) led to a low root:shoot (LRS) ratio, and environmental factors had a greater impact on AGB. For the LPUE pattern, soil water content (SWC), pH, and soil total nitrogen (STN) mainly affected the HRS ratio. Soil water content and STN influenced the HRS ratio through a positive effect on BGB . For the HPUE pattern, SWC, STN, and plant abundance (PA) predominantly regulate the LRS ratio, while biodiversity (plant abundance) affects the LRS ratio by positively affecting AGB. Our results highlight the differential impact of precipitation use efficiency on aboveground and belowground biomass allocation. This is important for monitoring the impact of drought events on plant biomass, improving productivity assessment models in arid and semi‐arid regions, and assessing local carbon storage accurately.

Phenotypic and transcriptomic responses of cultivated sunflower seedlings (Helianthus annuus L.) to four abiotic stresses

Plants encounter and respond to numerous abiotic stresses during their lifetimes. These stresses are often related and could therefore elicit related responses. There are, however, relatively few detailed comparisons between multiple different stresses at the molecular level. Here, we investigated the phenotypic and transcriptomic response of cultivated sunflower (Helianthus annuus L.) seedlings to three water-related stresses (i.e., dry-down, an osmotic challenge, and salt stress), as well as a generalized low-nutrient stress. All four stresses negatively impacted seedling growth, with the nutrient stress having a more divergent response from control as compared to the water-related stresses. Phenotypic responses were consistent with expectations for growth in low-resource environments, including increased (i.e., less negative) carbon fractionation values and leaf C:N ratios, as well as increased belowground biomass allocation. The number of differentially expressed genes (DEGs) under stress was greater in leaf tissue, but roots exhibited a higher proportion of DEGs unique to individual stresses. Overall, the three water-related stresses had a more similar transcriptomic response to each other vs. nutrient stress, though this pattern was more pronounced in root vs. leaf tissue. In contrast to our DEG analyses, co-expression network analysis revealed that there was little indication of a shared response between the four stresses in despite the majority of DEGs being shared between multiple stresses. Importantly, osmotic stress, which is often used to simulate drought stress in experimental settings, had little transcriptomic resemblance to true water limitation (i.e., dry-down) in our study, calling into question its utility as a means for simulating drought.

Precipitation-mediated responses of plant biomass production and allocation to changing soil pH in semiarid grasslands

Biomass and its allocation are crucial to understand carbon cycling. Climate change and human disturbance potentially alter soil pH, but it remains unclear how biomass production and its partitioning respond to altered soil pH. Here, we investigated how changing soil pH (acidification and alkalization) affected above- and belowground biomass production and allocation (hereafter AB, BB and BB/AB, respectively) at nine sites representing desert, typical, and meadow grasslands with increasing precipitation across a 1500-km transect in Inner Mongolia, China. The results showed that soil acidification reduced AB in typical and meadow grasslands, whereas soil alkalization reduced AB in desert and typical grasslands. Changing soil pH did not affect BB in desert or typical grasslands, but reduced BB in meadow grasslands. Altered soil pH negatively impacted species richness, Shannon-Wiener diversity, and dominant species biomass across all grasslands. BB/AB was increased by soil acidification and alkalization along the study transect. Soil C, N and C:N ratio were not affected by changing soil pH in three types of grasslands. Response ratios of AB, BB/AB and dominant species biomass were positively correlated with precipitation under altered soil pH, whereas response ratios of BB were negatively correlated with precipitation under soil alkalization. Variance partitioning analysis showed that the response of biomass and its allocation to altered soil pH were mainly explained by community composition but not soil fertility. Structural equation model analysis demonstrated that precipitation mediated the responses of biomass distribution to soil pH changes in semi-arid grasslands mainly via altering the response of dominant species. Our results indicate that dry grasslands with low productivity and precipitation generally allocate more C to belowground than wet grasslands regardless of pH change. These findings can improve our predictions of carbon dynamics in grassland ecosystems under climate change.

Plant functional types drive differential responses of grassland ecosystem functions along a precipitation gradient

Climatic variations can profoundly impact the functioning of grassland ecosystem, and understanding the drivers is critical for predicting ecosystem function under future climate. However, whether and how the responses of multiple grassland ecosystem functions to precipitation across spatial scales are controlled by the differential responses of the dominant plant functional types (PFT) remains unclear, mainly due to the scarcity of regional observational data. Here, we examined grassland ecosystem functions (aboveground plant biomass, belowground plant biomass, soil organic carbon, and total nitrogen) with different dominant plant functional types across 100 sites of four precipitation gradients in the Northeastern Qinghai-Tibetan Plateau by the moisture index. Across the broad precipitation gradient, higher mean annual precipitation significantly increased the plant species diversity and grassland ecosystem functions (aboveground plant biomass, soil organic C, and total N). From arid zone to humid zone, the above- and below-ground plant biomass allocation shifted to aboveground, and the trade-off between above- and below-ground plant biomass was the largest in semi-humid zone (0.13). Moreover, above- and below-ground plant biomass in arid, semi-arid, and semi-humid zones were driven by the grass-dominated communities (52%∼91%), while the humid zone was driven by the sedge- and forb- dominated communities (36%∼46%). The results of structural equational models further showed that the effects of precipitation on the ecosystem functions (soil organic carbon and total nitrogen) were mediated by either aboveground plant biomass (mainly in sedge-dominated communities) or species diversity (forb- and grass-dominated communities). These findings reveal that the asynchrony responses of different PFT-dominated communities to precipitation are altering grassland ecosystem functions, which provides the novel insight that plant functional group is particularly important for predicting ecosystem functioning under future climate conditions.

Simulated hurricane-induced changes in light and nutrient regimes change seedling performance in Everglades forest-dominant species.

Wind damage from cyclones can devastate the forest canopy, altering environmental conditions in the understory that affect seedling growth and plant community regeneration. To investigate the impact of hurricane‐induced increases in light and soil nutrients as a result of canopy defoliation, we conducted a two‐way factorial light and nutrient manipulation in a shadehouse experiment. We measured seedling growth of the dominant canopy species in the four Everglades forest communities: pine rocklands (Pinus elliottii var densa), cypress domes (Taxodium distichum), hardwood hammocks, and tree islands (Quercus virginiana and Bursera simaruba). Light levels were full sun and 50% shade, and nutrient levels coupled with an additional set of individuals that were subjected to a treatment mimicking the sudden effects of canopy opening from hurricane‐induced defoliation and the corresponding nutrient pulse. Seedlings were measured weekly for height growth and photosynthesis, with seedlings being harvested after 16 weeks for biomass, leaf area, and leaf tissue N and 13C isotope ratio. Growth rates and biomass accumulation responded more to differences in soil nutrients than differences in light availability, with largest individuals being in the high nutrient treatments. For B. simaruba and P. elliottii, the highest photosynthetic rates occurred in the high light, high nutrient treatment, while T. distichum and Q. virginiana photosynthetic rates were highest in low light, high nutrient treatment. Tissue biomass allocation patterns remained similar across treatments, except for Q. virginiana, which altered above‐ and belowground biomass allocation to increase capture of limiting soil and light resources. In response to the hurricane simulation treatment, height growth increased rapidly for Q. virginiana and B. simaruba, with nonsignificant increases for the other two species. We show here that ultimately, hurricane‐adapted, tropical species may be more likely to recolonize the forest canopy following a large‐scale hurricane disturbance.

Above- and belowground biomass allocation and its regulation by plant density in six common grassland species in China.

Above- and belowground biomass allocation is an essential plant functional trait that reflects plant survival strategies and affects belowground carbon pool estimation in grasslands. However, due to the difficulty of distinguishing living and dead roots, estimation of biomass allocation from field-based studies currently show large uncertainties. In addition, the dependence of biomass allocation on plant species, functional type as well as plant density remains poorly addressed. Here, we conducted greenhouse manipulation experiments to study above- and belowground biomass allocation and its density regulation for six common grassland species with different functional types (i.e., C3 vs C4; annuals vs perennials) from temperate China. To explore the density regulation on the biomass allocation, we used five density levels: 25, 100, 225, 400, and 625 plant m-2. We found that mean root to shoot ratio (R/S) values ranged from 0.04 to 0.92 across the six species, much lower than those obtained in previous field studies. We also found much lower R/S values in annuals than in perennials (C. glaucum and S. viridis vs C. squarrosa, L. chinensis, M. sativa and S. grandis) and in C4 plants than in C3 plants (C. squarrosa vs L. chinensis, M. sativa and S. grandis). In addition to S. grandis, plant density had significant effects on the shoot and root biomass fraction and R/S for the other five species. Plant density also affected the allometric relationships between above- and belowground biomass significantly. Our results suggest that R/S values obtained from field investigations may be severely overestimated and that R/S values vary largely across species with different functional types. Our findings provide novel insights into approximating the difficult-to-measure belowground living biomass in grasslands, and highlight that species composition and intraspecific competition will regulate belowground carbon estimation.

Belowground net primary productivity and biomass allocation in response to different restoration measures in a salt-alkali-degraded Songnen meadow

The understanding of belowground productivity restoration and biomass allocation is especially important for the restoration of salt-alkali-degraded meadows. Despite this, it remains unclear how belowground net primary productivity (BNPP) and biomass allocation respond to combined restoration measures. To assess the effects of different combined restoration measures on BNPP and biomass allocation, a field experiment was carried out in a salt-alkali-degraded meadow from 2012 to 2014. The restoration treatments included: free grazing (FG), enclosure with no-ploughing (E), enclosure with ploughing alone (EP), enclosure with ploughing and N addition (EPF), enclosure with ploughing and mulching (EPM), and enclosure with ploughing plus both N addition and mulching (EPFM). Our results across all three years showed that relative to FG, all the other restoration measures, whether single or combined, significantly increased aboveground net primary productivity (ANPP), BNPP and net primary productivity (NPP). The response of BNPP to different restoration measures was stronger than the ANPP response, resulting in a higher root fraction. The trends were clearer in the EPF, EPM and EPFM treatments, suggesting that more C was allocated to belowground roots with N addition and mulching. The strong increases in NPP and BNPP and belowground biomass allocation in the EPF, EPM and EPFM treatments were mainly due to significant increases in the density of important species (e.g. Chloris virgata and Puccinellia tenuiflora) and soil moisture, and strong decreases in soil pH and EC. Our findings demonstrate that vegetation cover, species richness and productivity (ANPP + BNPP) should be considered as important indicators when evaluating the recovery of severely degraded grassland. Our results further suggest that the combined measures, including enclosure and ploughing plus N addition and mulching, are effective ways to accelerate restoration in salt-alkali-degraded meadows.