Above‐ and below‐ground plant biomass response to experimental warming in northern Alaska

Question: Does experimental warming, designed to simulate future warming of the Arctic, change the biomass allocation and mycorrhizal infection of tundra plants? Location: High Arctic tundra near Barrow, Alaska, USA (71°18′N 156°40′W). Methods: Above and below ground plant biomass of all species was harvested following 3–4 yr of 1‐2°C of experimental warming. Biomass allocation and arbuscular mycorrhizal infection were also examined in the two dominant species, Salix rotundifolia and Carex aquatilis. Results: Above‐ground biomass of graminoids increased in response to warming but there was no difference in total plant biomass or the ratio of above‐ground to below‐ground biomass for the community as a whole. Carex aquatilis increased above‐ground biomass and proportionally allocated more biomass above ground in response to warming. Salix rotundifolia increased the amount of above‐ and below‐ground biomass allocated per leaf in response to warming. Mycorrhizal infection rates showed no direct response to warming, but total abundance was estimated to have likely increased in response to warming owing to increased root biomass of S. rotundifolia. Conclusions: The community as a whole was resistant to short‐term warming and showed no significant changes in above‐ or below‐ground biomass despite significant increases in above‐ground biomass of graminoids. However, the patterns of biomass allocation for C. aquatilis and S. rotundifolia did change with warming. This suggests that long‐term warming may result in changes in the above‐ground to below‐ground biomass ratio of the community.     

Below‐ground biomass and productivity of a grazed site and a neighbouring ungrazed exclosure in a grassland in central Argentina

Abstract We estimated the below‐ground net plant productivity (BNPP) of different biomass components in an intensively and continuously 45‐ha grazed site and in a neighbouring exclosure ungrazed for 16 years for a natural mountain grassland in central Argentina. We measured approximately twice as much dead below‐ground biomass in the grazed site as in the ungrazed site, with a strong concentration of total below‐ground biomass towards the upper 10 cm of the soil layer in both sites. The main contribution to total live biomass was accounted for by very fine (<0.5 mm) and fine roots (0.5–1.0 mm) both at the grazed (79%) and at the ungrazed (81%) sites. We measured more dead biomass for almost all root components, more live biomass of rhizomes, tap roots and bulbs, and less live biomass of thicker roots (>1 mm) in the grazed site. The seasonal variation of total live below‐ground biomass mainly reflected climate, with the growing season being limited to the warmer and wetter portion of the year, but such variation was higher in the grazed site. Using different methods of estimation of BNPP, we estimated maximum values of 1241 and 723 g m^?2 year^?1 for the grazed and ungrazed sites, respectively. We estimated that very fine root productivity was almost twice as high at the grazed site as at the ungrazed one, despite the fact that both sites had similar total live biomass, and root turnover rate was twofold at the grazed site.     

Factors determining plant–neighbour interactions on different spatial scales in young species-rich grassland communities

In naturally colonised species-rich grassland communities, we examined the properties of a plant’s aboveground neighbourhood that affect its performance (aboveground biomass). To this end a range of neighbourhood parameters were measured: number, biomass and species richness of the neighbours, number and biomass of the conspecific neighbours, and light availability at the base of the target plant. We also determined at which neighbourhood size the strongest target plant–neighbour interactions occurred, and whether conspecific neighbours affected competitively stronger or weaker target species differently. Target plant performance varied with target identity, and was significantly affected by light availability and the number of neighbouring plants (neighbourhood density). Depending on the target species, there was also an effect of total neighbour biomass on plant performance. The target plants were most strongly affected by their neighbours within a 3-cm distance, which could account for 78% of the variance in target biomass. Number or biomass of the conspecific neighbours did not contribute to the explanation of target performance in any of the target species. Whereas in an 8-cm neighbourhood the amount of light penetration was the strongest predictor of target performance, the number of neighbours was more important in a 3-cm neighbourhood. These experimental results might be useful to extend existing neighbourhood competition models for one or two species to multi-species competition models.     

Effects of a nematode Meloidogyne incognita and its interaction with above‐ground herbivory on an invasive wetland plant,alligator weed (Alternanthera philoxeroides)

Invasive plants may be attacked both above ground and below ground. Few studies have, however, investigated the simultaneous effects of above‐ground and below‐ground herbivory. In the present study, we report the effects of beetle herbivory and nematode infection on alligator weed, Alternanthera philoxeroides, an invasive plant in China. We found that the root‐knot nematode Meloidogyne incognita widely occurred on the plant in south China. To examine its effect on the plant in conjunction with above‐ground herbivory, we conducted a field common garden experiment with a local insect defoliator, Cassida piperata. We also included the native congener Alternanthera sessilis in our experiments for a comparison of the response of invasive and native species. We found no significant effects on plant biomass of the nematode infection in conjunction with the above‐ground herbivory. Further chemical analysis, however, showed that the water‐soluble carbohydrate content in roots of A. philoxeroides was significantly increased in plants attacked by both the nematode and the herbivore compared with the water‐soluble carbohydrate content in plants attacked by only the nematode or herbivore alone. We found no such change in the native congener A. sessilis. Together these results may suggest that A. philoxeroides tolerates joint above‐ground and below‐ground damage by allocating more resources to below‐ground material.     

Facilitation and interference in subalpine meadows of the central Caucasus

Abstract. We studied the effects of neighbours on the biomass of seven randomly chosen species in species‐rich sub‐alpine meadows in the central Caucasus Mountains by comparing the performance of plants with neighbours removed experimentally to that of paired plants with their neighbours left intact. In most cases the removal of neighbours led to significant increases in vegetative and total above‐ground biomass implying the species were limited by competition. However, the neighbour removal led also to an increased leaf wilting for target plants, as well as to strong decline in reproductive effort for some species. We hypothesise that competition may be the prevailing type of interaction in species‐rich sub‐alpine meadow communities, but competitive effects on vegetative production may be balanced, if not outweighed, by facilitation, at least for some species. Such a balance may enhance species coexistence in communities.     

Estimating root biomass in Salix viminalis × Salix schwerinii cultivar “Olof” using the electrical capacitance method

Abstract

Non‐destructive assessment of root systems is important in order to understand and optimise the potential of resource capture and allocation by the plant. We studied the relationships between electrical capacitance (EC) and the below‐ and above‐ground biomass of willows. Cuttings of Salix viminalis × Salix schwerinii cv. Olof were maintained in pots and root development was followed up using a portable capacitance meter over the course of 2.5 months. Pot observations were compared with excavation of two‐year‐old established trees. A strong significant linear relationship (R ² = 0.81, p < 0.001) was obtained between EC and root biomass (dry weight [DW]) for the pot experiment. EC also showed good correlations with stem and leaf biomass, as well as with stem height. In the excavated willow trees, there was a strong logarithmic relationship between EC and root biomass (R ² = 0.66, p < 0.001). These results suggest that EC is a good estimator of below‐ground biomass in willow and may become useful in screening varieties for differences in root biomass traits, especially in distinguishing below‐ground resource allocation at an early stage.     

Root seasonal pattern,spatial distribution,and C:N ratio of matgrass pasture (Nardus stricta L.) in the Lombardy Prealps

Abstract

The aim of the present study was to investigate carbon and nutrient cycling and the role of root dynamics in terrestrial ecosystems such as large abandoned pastures and natural grasslands present in the Prealps, for which below‐ground processes are currently enigmatic. In particular, we quantified root/leaf biomass and C:N ratio throughout two growing seasons. Additionally, root traits such as root length density (RLD), root mass density (RMD), and root diameter classes (RDC) were also investigated with the aim of understanding the spatial distribution of roots in the soil. In our samples, we found that the roots could be divided into three main diameter classes and hence quantified the presence of each class along the soil profile. With regard to total root biomass, we found the occurrence of two peaks of biomass accumulation during the growth season, and when biomass accumulation was compared with climatic data, it was impossible to obtain a clear indication of the root turnover rate. In fact, the strong influence of grazing on the above‐ground biomass could have affected, in turn, root biomass. In future, this possible complication will be avoided by repeating the measurements within enclosures to avoid grazing interference. We found that C:N ratio remained constant, with a single peak, suggesting a lower root decomposition during the warmest period (August 2006). The concentration of nitrogen in roots decreased with depth as a result of a decrease in roots with smaller diameters. The reverse was found for carbon content, which increased with depth, probably due to an increase in roots with larger diameters. This study represents the first attempt to estimate root turnover rates in this prealpine ecosystem, which have been analysed to date only for the above‐ground biomass.     

Soil warming and CO2 enrichment induce biomass shifts in alpine tree line vegetation

Responses of alpine tree line ecosystems to increasing atmospheric CO₂ concentrations and global warming are poorly understood. We used an experiment at the Swiss tree line to investigate changes in vegetation biomass after 9 years of free air CO₂ enrichment (+200 ppm; 2001–2009) and 6 years of soil warming (+4 °C; 2007–2012). The study contained two key tree line species, Larix decidua and Pinus uncinata, both approximately 40 years old, growing in heath vegetation dominated by dwarf shrubs. In 2012, we harvested and measured biomass of all trees (including root systems), above‐ground understorey vegetation and fine roots. Overall, soil warming had clearer effects on plant biomass than CO₂ enrichment, and there were no interactive effects between treatments. Total plant biomass increased in warmed plots containing Pinus but not in those with Larix. This response was driven by changes in tree mass (+50%), which contributed an average of 84% (5.7 kg m^?2) of total plant mass. Pinus coarse root mass was especially enhanced by warming (+100%), yielding an increased root mass fraction. Elevated CO₂ led to an increased relative growth rate of Larix stem basal area but no change in the final biomass of either tree species. Total understorey above‐ground mass was not altered by soil warming or elevated CO₂. However, Vaccinium myrtillus mass increased with both treatments, graminoid mass declined with warming, and forb and nonvascular plant (moss and lichen) mass decreased with both treatments. Fine roots showed a substantial reduction under soil warming (?40% for all roots <2 mm in diameter at 0–20 cm soil depth) but no change with CO₂ enrichment. Our findings suggest that enhanced overall productivity and shifts in biomass allocation will occur at the tree line, particularly with global warming. However, individual species and functional groups will respond differently to these environmental changes, with consequences for ecosystem structure and functioning.     

Effects of physical connection and genetic identity of neighbouring ramets on root-placement patterns in two clonal species

Root-placement patterns were examined in the clonal species Glechoma hederacea and Fragaria vesca when grown with different types of neighbours. Three different patterns were predicted as consequences of different types of interactions between roots: the avoidance pattern if root growth decreases in the presence of neighbouring roots; the intrusive pattern if root growth increases towards neighbouring roots; and the unresponsive pattern if root growth is unaffected by neighbouring roots. Experiments were conducted in which physical connection between ramets, and the genetic identity of neighbouring ramets, were manipulated. The patterns of distribution of entire root systems and elongation rates of individual roots were measured. Root systems and individual roots of G. hederacea avoided contact with roots of neighbouring ramets, irrespective of connection to the neighbour and its genetic or specific identity. In contrast, F. vesca roots grew equally towards and away from intraspecific ramet neighbours and their elongation was stimulated by contact with roots of G. hederacea ramets. These results demonstrate that root-placement patterns of plants grown with different types of neighbours vary between species, and suggest that factors additional to resource depletion could be involved in their development.     

Old‐field grassland successional dynamics following cessation of chronic disturbance

Question: Does increasing Festuca canopy cover reduce plant species richness and, therefore, alter plant community composition and the relationship of litter to species richness in old‐field grassland? Location: Southeastern Oklahoma, USA. Methods: Canopy cover by species, species richness, and litter mass were collected within an old‐field grassland site on 16, 40 m × 40 m plots. Our study was conducted during the first three years of a long‐term study that investigated the effects of low‐level nitrogen enrichment and small mammal herbivory manipulations. Results: Succession was altered by an increase in abundance of Festuca over the 3‐yr study period. Species richness did not decline with litter accumulation. Instead, Festuca increased most on species‐poor plots, and Festuca abundance remained low on species‐rich plots. Conclusions: Festuca may act as an invasive transformer‐species in warm‐season dominated old‐field grasslands, a phenomenon associated more with invasions of cool‐season grasses at higher latitudes in North America.     

Semi‐polar root exudates in natural grassland communities

In the rhizosphere, plants are exposed to a multitude of different biotic and abiotic factors, to which they respond by exuding a wide range of secondary root metabolites. So far, it has been unknown to which degree root exudate composition is species‐specific and is affected by land use, the local impact and local neighborhood under field conditions. In this study, root exudates of 10 common grassland species were analyzed, each five of forbs and grasses, in the German Biodiversity Exploratories using a combined phytometer and untargeted liquid chromatography‐mass spectrometry (LC‐MS) approach. Redundancy analysis and hierarchical clustering revealed a large set of semi‐polar metabolites common to all species in addition to species‐specific metabolites. Chemical richness and exudate composition revealed that forbs, such as Plantago lanceolata and Galium species, exuded more species‐specific metabolites than grasses. Grasses instead were primarily affected by environmental conditions. In both forbs and grasses, plant functional traits had only a minor impact on plant root exudation patterns. Overall, our results demonstrate the feasibility of obtaining and untargeted profiling of semi‐polar metabolites under field condition and allow a deeper view in the exudation of plants in a natural grassland community.     

Temporal asynchrony in fine‐root biomass may contribute to shrub and grass coexistence in mixed patches

We described seasonal changes in fine‐root biomass of a grass and a shrub dominant species in a plant community characteristic of the arid Patagonian Monte and then we inferred to want extent the observed differences could contribute to the species coexistence. We selected representative plant patches of the natural vegetation arrangement consisting of one isolated plant of the dominant shrub Larrea divaricata (Ld), grass patches formed by one or more bunches of the dominant grass Nassella tenuis (Nt), and mixed patches consisting of one individual of L. divaricata with bunches of N. tenuis under its canopy (LdNt). We assessed the biomass and temporal changes in fine roots of each species in the upper soil (50 cm depth) of each patch type at three‐month intervals during 2 years. The temporal series of fine‐root biomass were compared among patch types and in relation to above‐ground phenology, as well as climate variables (precipitation, arid index and air temperature). Seasonal changes in fine‐root biomass showed similar cycles in the three plant patches with a maximum in spring. The maximum increase in root biomass in Ld and Nt patches occurred during the onset of reproductive growth in winter and spring, respectively. Fine‐root changes in LdNt patches mimicked that in Ld patches. Precipitation inputs were significantly positively and temperature negatively related to fine‐root changes in Nt patches. Fine‐root changes in Ld and LdNt patches were related to the aridity index (positively) and temperature (negatively). We concluded that the observed asynchronies in the date of the largest increases in root biomass and its climate control between the studied grass and shrub species could contribute to the coexistence of plants of both life forms when they overlap their root systems growing in mixed patches. Mechanisms underlying the root patterns observed should be further explored.     

Predicting competitive interactions between pioneer plant species by using plant traits

Abstract. A competitive effect hierarchy for 15 Namaqualand pioneer plant species was established by using the mean mass of the phytometer (Dimorphotheca sinuata) when grown in combination with itself and 14 other species. There were no clear groupings of species in the hierarchy. This competitive hierarchy (gradient) indicated which species are strong competitors (resulting in a low phytometer mass) with D. sinuata and which species are weak competitors (resulting in a high phytometer mass). Each plant species has a certain combination of plant traits which determines its life history strategy and competitive ability. Regressions of various plant traits (measured on plants grown singly) against phytometer biomass indicated which traits were significantly correlated. The traits, most being size-related, were: maximum shoot mass, total mass, stem mass, reproductive mass, leaf area, stem allocation, specific leaf area (SLA), vegetative height × diameter, leaf area ratio (LAR); and mean number of days to flower initiation. A forward stepwise multiple regression of the significant traits was used to determine an equation to predict competitive effect.     

Large‐scale pattern of biomass partitioning across China's grasslands

Aim To investigate large‐scale patterns of above‐ground and below‐ground biomass partitioning in grassland ecosystems and to test the isometric theory at the community level. Location Northern China, in diverse grassland types spanning temperate grasslands in arid and semi‐arid regions to alpine grasslands on the Tibetan Plateau. Methods We investigated above‐ground and below‐ground biomass in China's grasslands by conducting five consecutive sampling campaigns across the northern part of the country during 2001–05. We then documented the root : shoot ratio (R/S) and its relationship with climatic factors for China's grasslands. We further explored relationships between above‐ground and below‐ground biomass across different grassland types. Results Our results indicated that the overall R/S of China's grasslands was larger than the global average (6.3 vs. 3.7). The R/S for China's grasslands did not show any significant trend with either mean annual temperature or mean annual precipitation. Above‐ground biomass was nearly proportional to below‐ground biomass with a scaling exponent (the slope of log–log linear relationship between above‐ground and below‐ground biomass) of 1.02 across various grassland types. The slope did not differ significantly between temperate and alpine grasslands or between steppe and meadow. Main conclusions Our findings support the isometric theory of above‐ground and below‐ground biomass partitioning, and suggest that above‐ground biomass scales isometrically with below‐ground biomass at the community level.     

Seasonal storage of nutrients by perennial herbaceous species in undisturbed and disturbed deciduous hardwood forests

Question: Pollution and eutrophication of surface water is increasingly a problem in agricultural landscapes. Do intact (relatively undisturbed) and degraded forests differ in seasonal nutrient storage and therefore potential to ameliorate nutrient pollution? Location: United States, Midwestern region. Methods: We used three sets of paired plots, where intact plots were located close to disturbed woodlands. Herbaceous perennials located in eight 0.25 m² quadrats in the plots were harvested (in spring and mid‐summer), dried, separated into above‐ and below‐ground plant parts, and weighed to determine biomass. Nitrogen, phosphorus and potassium content of the plant tissues were then determined, and these data combined with biomass to estimate nutrient storage. Results: In spring, intact sites had 62% greater above‐ground biomass than disturbed sites and 75% greater below‐ground biomass. In summer, below‐ground biomass of intact plots was still much greater than that of disturbed plots (73 percent), but above‐ground biomass was similar. Nutrient tissue concentration generally did not differ, nor did soil nutrient levels. The disturbed sites were largely missing one group of species, the spring ephemerals, and this accounted for the difference in biomass and nutrient storage between sites. Conclusions: Relatively undisturbed woodlands in our study had a much greater capacity to store nutrients, and therefore ameliorate nutrient pollution, in early spring. This is significant because spring is also the time of highest potential leaching of nutrients into surface water.     

Light partitioning among species and species replacement in early successional grasslands

Abstract. We studied canopy structure, shoot architecture and light harvesting efficiencies of the species (photon flux captured per unit above‐ground plant mass) in a series of exclosures of different age (up to 4.5 yr) in originally heavily grazed grassland in N Japan.Vegetation height and Leaf Area Index (LAI) increased in the series and Zoysia japonica, the dominant in the beginning, was replaced by the much taller Miscanthus sinensis. We showed how this displacement in dominance can be explained by inherent constraints on the above‐ground architecture of these two species. In all stands light capture of plants increased with their above‐ground biomass but taller species were not necessarily more efficient in light harvesting. Some subordinate species grew disproportionally large leaf areas and persisted in the shady undergrowth. Some other species first grew taller and managed to stay in the better‐lit parts of the canopy, but ultimately failed to match the height growth of their neighbours in this early successional series. Their light harvesting efficiencies declined and this probably led to their exclusion. By contrast, species that maintained their position high in the canopy managed to persist in the vegetation despite their relatively low light harvesting efficiencies. In the tallest stands ‘later successional’ species had higher light harvesting efficiencies for the same plant height than ‘early successional’ species which was mostly the result of the greater area to mass ratio (specific leaf area, SLA) of their leaves. This shows how plant stature, plasticity in above‐ground biomass partitioning, and architectural constraints determine the ability of plants to efficiently capture light, which helps to explain species replacement in this early successional series.     

A regional‐scale consideration of the effects of species richness on above‐ground biomass in temperate natural grasslands of China

Question: Can species richness be a predictor for above‐ground biomass in natural grasslands at a regional scale? Location: A total of 647 sites across temperate natural grasslands of northern China. Methods: Structural equation modelling (SEM) was used to examine the effect of species richness on above‐ground biomass. Asymptotically distribution‐free estimation was selected for parameter estimation. The SEM process was performed at five sample sizes (n=50, 100, 200, 300 and 647). Spatial structure in the original data was examined by calculating Moran's I. Results: SEM run at n=647 revealed a positive effect of species richness on above‐ground biomass after controlling for the influences of bioclimatic factors and grazing. At the four reduced sample size levels, the positive effect held true for most cases in 400 observations (>92%). Conclusions: Contrary to observations in previous studies in natural grasslands, our data showed a positive effect of species richness on above‐ground biomass. This suggests that, as a short‐term effect of diversity on productivity, niche complementarity among coexisting species tends to be an important process in arid and semi‐arid natural grasslands. We hold that biodiversity conservation is among the fundamental approaches required to maintain productivity of grasslands in arid and semi‐arid areas.     

Root damage to apple plants by cockchafer larvae induces a change in volatile signals below‐ and above‐ground

Volatile organic compounds (VOCs) mediate communication between plants and insects. Plants under insect herbivore attack release VOCs either at the site of attack or systemically, indicating within‐plant communication. Some of these VOCs, which may be induced only upon herbivore attack, recruit parasitoids and predatory insects to feed on the attacking insects. Moreover, some plants are able to ‘eavesdrop’ on herbivore‐induced plant volatiles (HIPVs) to prime themselves against impending attack; such eavesdropping exemplifies plant–plant communication. In apple orchards, the beetle Melolontha melolontha L. (Coleoptera: Scarabaeidae) is an important insect pest whose larvae live and feed on roots for about 4 years. In this study, we investigated whether the feeding activity of M. melolontha larvae (1) alters the volatile profile of apple roots, (2) induces the release of HIPVs systemically in the leaves, and (3) whether infested plants communicate to neighbouring non‐infested conspecifics through HIPVs. To answer these questions, we collected constitutive VOCs from intact M9 roots as well as M. melolontha larvae‐damaged roots using a newly designed ‘rhizobox’, to collect root‐released volatiles in situ, without damaging the plant root system. We also collected VOCs from the leaf‐bearing shoots of M9 whose roots were under attack by M. melolontha larvae and from shoots of neighbouring non‐infested conspecifics. Gas chromatography‐mass spectrometry analysis showed that feeding activity of M. melolontha larvae induces the release of specific HIPVs; for instance, camphor was found in the roots only after larvae caused root damage. Melolontha melolontha also induced the systemic release of methyl salicylate and (E,E)‐α‐farnesene from the leaf‐bearing shoots. Methyl salicylate and (E,E)‐α‐farnesene were also released by the shoots of non‐infested neighbouring conspecifics. These phenomena indicate the induction of specific VOCs below‐ and above‐ground upon M. melolontha larvae feeding on apple roots as well as plant–plant communication in apple plants.     

Resource competition and fire‐regulated nutrient demand in carnivorous plants of wet pine savannas

Question: What are the mechanisms by which fire reduces competition for both a short‐lived and a long‐lived species in old‐growth ground‐cover plant communities of wet pine savannas (originally Pinus palustris, replaced by P. elliottii)? Location: Outer coastal plain of southeastern Mississippi, USA. Methods: I reviewed previous competition experiments and proposed a new hypothesis to explain the relationship between fire, competition, and species co‐existence in wet longleaf pine savannas. The first study is about growth and seedling emergence responses of a short‐lived carnivorous plant, Drosera capillaris, to reduction in below‐ground competition and above‐ plus below‐ground competition. The second study deals with growth and survival responses of a long‐lived perennial carnivorous plant, Sarracenia alata, to neighbour removal and prey‐exclusion to determine if a reduction in nutrient supply increased the intensity of competition in this nutrient‐poor system. Results: Fire increased seedling emergence of the short‐lived species by reducing above‐ground competition through the destruction of above‐ground parts of plants and the combustion of associated litter. Prey exclusion did not increase competitive effects of neighbours on the long‐lived species. However, because the experiment was conducted in a year without fire, shade reduced nutrient demand, which may have obviated competition for soil nutrients between Sarracenia alata and its neighbours. Conclusion: Repeated fires likely interact with interspecific differences in nutrient uptake to simultaneously reduce both above‐ground competition and competition for nutrients in old‐growth ground cover communities in pine savannas. Restoration practitioners should consider the possibility that the composition of the plant community is just as important as fire in ensuring that frequent fires maintain species diversity.