Below-ground herbivory promotes soil nutrient transfer and root growth in grassland

Extremely little is known about the ecosystem-level implications of below-ground herbivory, which often represents the dominant form of consumption of primary productivity. We provide the first empirical evidence that low levels of below-ground herbivory may promote soil nutrient flux and root growth of both host plants and companion plants. Low levels of white clover ( Trifolium repens L.) root infection by clover cyst nematodes ( Heterodera trifolii Goffart) increased root growth by 141% and 219% in the host plant and the uninfected neighbouring grass ( Lolium perenne L.), respectively. Root infection increased the size of the soil microbial biomass in the root zone and the transfer of ¹⁵N from the host plant to soil and the neighbouring grass. These data suggest that low amounts of below-ground herbivory may increase the transfer of plant carbon and nitrogen below-ground, leading to increases in root growth and soil nutrient recycling in grasslands. Presumably, such interactions will influence the competitive interactions between plant species, altering plant community structure in grasslands.     

Effect of soil nutrient heterogeneity on the symmetry of belowground competition

Size variability in plant populations has been extensively studied andmuch of this inquiry has focused on the types of competition that are involvedin increasing or decreasing size inequality. It is often assumed thatcompetition for light is size-asymmetric, meaning that a plant canpotentially dominate a competitive relationship through shading if it is tallerthan its competitor. The light resources obtained by the taller plant are thusdisproportionate to its size. In contrast, competition for soil resources maybemore size-symmetric, the amount of soil nutrients obtained seems to be indirect proportion to a plant's size. Most studies examining belowgroundcompetition have used homogeneouslydistributed nutrient resources. Soil homogeneity could makesize-asymmetric belowground competition unlikely, but homogeneity is notoften found in nature. In this study I use a greenhouse pot experimentutilizingIpomoea tricolor to examine the hypothesis thatsize-asymmetric competition for soil nutrients may result when soilresource distribution is spatially heterogeneous. The results did not supportthe hypothesis of belowgroundsize-asymmetric competition. Differences between experimental treatmentsand controls were not statistically significant suggesting size-symmetriccompetition. Implications of these results are discussed.     

植物根系养分捕获塑性与根竞争

为了更有效地从土壤中获取养分, 植物根系在长期的进化与适应中产生了一系列塑性反应, 以响应自然界中广泛存在的时空异质性。同时, 植物根系的养分吸收也要面对来自种内和种间的竞争。多种因素都会影响植物根竞争的结果, 包括养分条件、养分异质性的程度、根系塑性的表达等。竞争会改变植物根系的塑性反应, 比如影响植物根系的空间分布; 植物根系塑性程度差异也会影响竞争。已有研究发现根系具有高形态塑性和高生理塑性的植物在长期竞争过程中会占据优势。由于不同物种根系塑性的差异, 固定的对待竞争的反应模式在植物根系中可能并不存在, 其响应随竞争物种以及土壤环境因素的变化而变化。此外, 随着时间变化, 根系塑性的反应及其重要性也会随之改变。植物对竞争的反应可能与竞争个体之间的亲缘关系有关, 有研究表明亲缘关系近的植物可能倾向于减小彼此之间的竞争。根竞争对植物的生存非常重要, 但目前还没有研究综合考虑植物的各种塑性在根竞争中的作用。另外根竞争对群落结构的影响尚待深入的研究。     

Interactive effects of soil fertility and herbivory on Brassica nigra

Soil nutrient availability may affect both the amount of damage that plants receive from herbivores and the ability of plants to recover from herbivory, but these two factors are rarely considered together. In the experiment reported here, I examined how soil fertility influenced both the degree of defoliation and compensation for herbivory for Brassica nigra plants damaged by Pieris rapae caterpillars. Realistic levels of defoliation were obtained by placing caterpillars on potted host plants early in the life cycle and allowing them to feed until just before pupation on the designated plant. Percent defoliation was more than twice as great at low soil fertility compared to high (48.2% and 21.0%, respectively), even though plants grown at high soil fertility lost a greater absolute amount of leaf area (38.2 cm² and 22.1 cm², respectively). At both low and high soil fertility, total seed number and mean mass per seed of damaged plants were equivalent to those of undamaged plants. Thus soil fertility did not influence plant compensation in terms of maternal fitness. However, the pathways used to achieve compensation in seed production were different at low and high soil fertility. At low soil fertility, relative leaf growth rates (area added per inital area per day) of damaged plants were drastically reduced over the second week of caterpillar feeding. Damaged plants recovered the leaf area lost to herbivory in the two weeks following insect removal by increasing leaf relative growth rates above the levels seen for undamaged plants, but the replacement of leaf tissue lost to herbivory came at the expense of stem biomass. At high soil fertility, relative leaf growth rates of damaged plants were similar to those of undamaged plants both over the second week of caterpillar feeding and following caterpillar removal, and stem biomass was not affected by herbivory. These results suggest that higher levels of soil nutrients increased the ability of plants to stay ahead of their herbivores as they were being eaten. Because damaged plants at high soil fertility were able to maintain leaf growth rates to a greater extent than damaged plants at low soil fertility, they did not fall as far behind undamaged plants over the period of insect feeding and did not have as much catching up to do after feeding ended to compensate for herbivory.     

Effects of aboveground herbivory on plants with long-term belowground biomass storage

Plant tolerance to herbivory is contingent on multiple traits and adaptive mechanisms, which makes it a complex response with ecological implications. In plants with long-term belowground storage, allocation of biomass to inaccessible parts belowground in response to folivory is a well-recognized tolerance mechanism. In temperate regions, spring growth from buried rootstock is common among winter deciduous plants and is often followed by regrowth after defoliation, both of which draws resources from the stored reserves. We developed a mathematical model to analyze this tolerance response in a winter deciduous plant with long-term belowground biomass when it is defoliated by a specialist insect folivore. The model explores how three closely associated traits—(1) belowground biomass allocation to roots, (2) spring utilization of stored reserves, and (3) post-defoliation regrowth capacity—modulate the persistence and dynamics of the plant and herbivore populations. Model results show that allocation to belowground storage is not only a critical component of tolerance but also influences the herbivore population dynamics in ways that depend on how and when plant biomass is allocated and used. Low belowground biomass allocation and high storage utilization combined with poor photosynthetic growth caused extirpation of the plant population by the defoliating insects. Stable coexistence of the plant at low biomass along with its specialist insect required a moderate amount of post-herbivory belowground allocation. High values of belowground biomass allocation, storage utilization, and photosynthetic growth resulted in sustained cycles of the herbivore and plant populations. Interestingly, utilization of stored reserves had conflicting influence on above and belowground biomass, and strongly affected herbivore population dynamics. Our model thus highlights the complexity of tolerance response when it involves multiple traits and mechanisms as evinced by winter deciduous plants. We close by discussing the implications of our findings for the contributions of defoliating insects to biocontrol programs.     

Community-aggregated plant traits interact with soil nutrient heterogeneity to determine ecosystem functioning

Background and aims

Spatial distribution of soil nutrients (soil heterogeneity) and availability have strong effects on above- and belowground plant functional traits. Although there is ample evidence on the tight links between functional traits and ecosystem functioning, the role played by soil heterogeneity and availability as modulators of such relationship is poorly known.

Methods

We conducted a factorial experiment in microcosms containing grasses, legumes and non-legume forbs communities differing in composition to evaluate how soil heterogeneity and availability (50 and 100 mg N) affect the links between traits and ecosystem functioning. Community-aggregated specific leaf area (SLA_agg) and specific root length (SRL_agg) were measured as both relevant response traits to soil heterogeneity and availability, and significant effect traits affecting ecosystem functioning (i.e., belowground biomass, β-glucosidase and acid phosphatase activities, and in situ N availability rate).

Results

SRL_agg was negatively and significantly associated to β-glucosidase, phosphatase and N availability rate in the high nutrient availability and heterogeneous distribution scenario. We found a significant negative relationship between SLA_agg and availability rate of mineral-N under low nutrient availability conditions.

Conclusions

Soil heterogeneity modulated the effects of both traits and nutrient availability on ecosystem functioning. Specific root length was the key trait associated with soil nutrient cycling and belowground biomass in contrasted heterogeneous soil conditions. The inclusion of soil heterogeneity into the trait-based response-effect framework may help to scale from plant communities to the ecosystem level.     

Sequential effects of root and foliar herbivory on aboveground and belowground induced plant defense responses and insect performance

Plants are often simultaneously or sequentially attacked by multiple herbivores and changes in host plants induced by one herbivore can influence the performance of other herbivores. We examined how sequential feeding on the plant Plantago lanceolata by the aboveground herbivore Spodoptera exigua and the belowground herbivore Agriotes lineatus influences plant defense and the performance of both insects. Belowground herbivory caused a reduction in the food consumption by the aboveground herbivore independent of whether it was initiated before, at the same time, or after that of the aboveground herbivore. By contrast, aboveground herbivory did not significantly affect belowground herbivore performance, but significantly reduced the performance of later arriving aboveground conspecifics. Interestingly, belowground herbivores negated negative effects of aboveground herbivores on consumption efficiency of their later arriving conspecifics, but only if the belowground herbivores were introduced simultaneously with the early arriving aboveground herbivores. Aboveground–belowground interactions could only partly be explained by induced changes in an important class of defense compounds, iridoid glycosides (IGs). Belowground herbivory caused a reduction in IGs in roots without affecting shoot levels, while aboveground herbivory increased IG levels in roots in the short term (4 days) but only in the shoots in the longer term (17 days). We conclude that the sequence of aboveground and belowground herbivory is important in interactions between aboveground and belowground herbivores and that knowledge on the timing of exposure is essential to predict outcomes of aboveground–belowground interactions.     

Consequences of insect herbivory on grape fine root systems with different growth rates

Herbivory tolerance has been linked to plant growth rate where plants with fast growth rates are hypothesized to be more tolerant of herbivory than slower-growing plants. Evidence supporting this theory has been taken primarily from observations of aboveground organs but rarely from roots. Grapevines differing in overall rates of new root production, were studied in Napa Valley, California over two growing seasons in an established vineyard infested with the sucking insect, grape phylloxera (Daktulosphaira vitifoliae Fitch). The experimental vineyard allowed for the comparison of two root systems that differed in rates of new root tip production (a 'fast grower', Vitis berlandieri x Vitis rupestris cv. 1103P, and a slower-growing stock, Vitis riparia x Vitis rupestris cv. 101-14 Mgt). Each root system was grafted with a genetically identical shoot system (Vitis vinifera cv. Merlot). Using minirhizotrons, we did not observe any evidence of spatial or temporal avoidance of insect populations by root growth. Insect infestations were abundant throughout the soil profile, and seasonal peaks in phylloxera populations generally closely followed peaks in new root production. Our data supported the hypothesis that insect infestation was proportional to the number of growing tips, as indicated by similar per cent infestation in spite of a threefold difference in root tip production. In addition, infested roots of the fast-growing rootstock exhibited somewhat shorter median lifespans (60 d) than the slower-growing rootstock (85 d). Lifespans of uninfested roots were similar for the two rootstocks (200 d). As a consequence of greater root mortality of younger roots, infested root populations in the fast-growing rootstock had an older age structure. While there does not seem to be a trade-off between potential growth rate and relative rate of root infestation in these cultivars, our study indicates that a fast-growing root system may more readily shed infested roots that are presumably less effective in water and nutrient uptake. Thus, differences in root tip production may be linked to differences in the way plants cope with roots that are infested by sucking insects.     

Interactive effects of soil temperature, atmospheric carbon dioxide and soil N on root development, biomass and nutrient uptake of winter wheat during vegetative growth

Nutrient requirements for plant growth are expected to rise in response to the predicted changes in CO(2) and temperature. In this context, little attention has been paid to the effects of soil temperature, which limits plant growth at early stages in temperate regions. A factorial growth-room experiment was conducted with winter wheat, varying soil temperature (10 degrees C and 15 degrees C), atmospheric CO(2) concentration (360 and 700 ppm), and N supply (low and high). The hypothesis was that soil temperature would modify root development, biomass allocation and nutrient uptake during vegetative growth and that its effects would interact with atmospheric CO(2) and N availability. Soil temperature effects were confirmed for most of the variables measured and 3-factor interactions were observed for root development, plant biomass components, N-use efficiency, and shoot P content. Importantly, the soil temperature effects were manifest in the absence of any change in air temperature. Changes in root development, nutrient uptake and nutrient-use efficiencies were interpreted as counterbalancing mechanisms for meeting nutrient requirements for plant growth in each situation. Most variables responded to an increase in resource availability in the order: N supply >soil temperature >CO(2).     

Interactive effects of plant-available soil silicon and herbivory on competition between two grass species

Background and Aims

The herbivore defence system of true grasses (Poaceae) is predominantly based on silicon that is taken up from the soil and deposited in the leaves in the form of abrasive phytoliths. Silicon uptake mechanisms can be both passive and active, with the latter suggesting that there is an energetic cost to silicon uptake. This study assessed the effects of plant-available soil silicon and herbivory on the competitive interactions between the grasses Poa annua, a species that has previously been reported to accumulate only small amounts of silicon, and Lolium perenne, a high silicon accumulator.

Methods

Plants were grown in mono- and mixed cultures under greenhouse conditions. Plant-available soil silicon levels were manipulated by adding silicon to the soil in the form of sodium silicate. Subsets of mixed culture pots were exposed to above-ground herbivory by desert locusts (Schistocerca gregaria).

Key Results

In the absence of herbivory, silicon addition increased biomass of P. annua but decreased biomass of L. perenne. Silicon addition increased foliar silicon concentrations of both grass species >4-fold. Under low soil-silicon availability the herbivores removed more leaf biomass from L. perenne than from P. annua, whereas under high silicon availability the reverse was true. Consequently, herbivory shifted the competitive balance between the two grass species, with the outcome depending on the availability of soil silicon.

Conclusions

It is concluded that a complex interplay between herbivore abundance, growth–defence trade-offs and the availability of soil silicon in the grasses'' local environment affects the outcome of inter-specific competition, and so has the potential to impact on plant community structure.     

Root growth of wetland plants with different root types

A wastewater culture system was designed to study the root growth of eight species of wetland plants with two different root types. The system included a plastic barrel for holding the wastewater and a foam plate for holding the plant. The results indicated that the root growth of the plants with fibril roots was faster than that of the plants with rhizomatic roots. The species with fibril roots had higher root number (1349 per plant) than species with rhizomatic roots (549 per plant) after ten weeks of cultivation. The average root biomass of plants with fibril roots was 11.3 g per plant, whereas that of plants with rhizomatic roots was 7.4 g per plant. Fine root biomass of diameter ≤ 1 mm constituted 51.9% of the total root biomass in plants with fibril roots, whereas it accounted for only 25.1% in plants with rhizomatic roots. The root surface area of the plants with fibril roots (6933 cm² per plant) was markedly larger than that of the species with rhizomatic roots (1897 cm² per plant). The species with rhizomatic roots showed a longer root lifespan (46.6 days) than those with fibril roots (34.8 days).     

两种不同根系类型湿地植物的根系生长

实验设计了一个水培系统,利用生活污水培养,对4种“须根型”植物美人蕉、风车草、象草和香根草和4种根茎型植物菖蒲、水鬼蕉、芦苇和水烛的根系生长进行比较研究。该系统由用于盛污水的塑料桶（顶部直径36．5cm,底部直径30．Ocm,高34．5cm）和用于固定植物于水面的泡沫板构成。每桶种植1株植物,每种种5株。水培至10周时,须根型植物的平均根数达到1349条／株,而根茎型植物的平均根数只有549条／株。实验结束（水培第21周）时,须根型植物的平均根生物量为11．3g／株,根茎型植物的平均根生物量为7．4g／株。须根型植物根系中,d〈1mm的细根生物量占根系总生物量的51．9％,而根茎型植物d〈1mm的细根的生物量只占25．1％。根茎型植物的根生物量与地上生物量的比值为0．2,显著高于须根型湿地植物（0．1）。须根型湿地植物的根系表面积（6933cm^2／株）极显著地高于根茎型湿地植物（1897cm^2／株）。根茎型湿地植物根的平均寿命（46．6d）较须根型湿地植物根的平均寿命（34．8d）长。美人蕉的平均根数达1871条／株,根表面积达到22832cm^2／株,远较其他种高。     

The effects of environmental heterogeneity on root growth and root/shoot partitioning

AIMS: The purpose of this Botanical Briefing is to stimulate reappraisal of root growth, root/shoot partitioning, and analysis of other aspects of plant growth under heterogeneous conditions. SCOPE: Until recently, most knowledge of plant growth was based upon experimental studies carried out under homogeneous conditions. Natural environments are heterogeneous at scales relevant to plants and in forms to which they can respond. Responses to environmental heterogeneity are often localized rather than plant-wide, and not always predictable from traditional optimization arguments or from knowledge of the ontogenetic trends of plants growing under homogeneous conditions. These responses can have substantial impacts, both locally and plant-wide, on patterns of resource allocation, and significant effects on whole-plant growth. Results from recent studies are presented to illustrate responses of plants, plant populations and plant communities to nutritionally heterogeneous conditions. CONCLUSIONS: Environmental heterogeneity is a constant presence in the natural world that significantly influences plant behaviour at a variety of levels of complexity. Failure to understand its effects on plants prevents us from fully exploiting aspects of plant behaviour that are only revealed under patchy conditions. More effort should be invested into analysis of the behaviour of plants under heterogeneous conditions.     

Interactive effects of soil temperature and moisture on Concord grape root respiration

Root respiration has important implications for understanding plant growth as well as terrestrial carbon flux with a changing climate. Although soil temperature and soil moisture often interact, rarely have these interactions on root respiration been studied. This report is on the individual and combined effects of soil moisture and temperature on respiratory responses of single branch roots of 1-year-old Concord grape (Vitis labruscana Bailey) vines grown in a greenhouse. Under moist soil conditions, root respiration increased exponentially to short-term (1 h) increases in temperature between 10 degrees C and 33 degrees C. Negligible increases in root respiration occurred between 33 degrees C and 38 degrees C. By contrast to a slowly decreasing Q10 from short-term temperature increases, when roots were exposed to constant temperatures for 3 d, the respiratory Q10 between 10 degrees C and 30 degrees C diminished steeply with an increase in temperature. Above 30 degrees C, respiration declined with an increase in temperature. Membrane leakage was 89-98% higher and nitrogen concentration was about 18% lower for roots exposed to 35 degrees C for 3 d than for those exposed to 25 degrees C and 15 degrees C. There was a strong interaction of respiration with a combination of elevated temperature and soil drying. At low soil temperatures (10 degrees C), respiration was little influenced by soil drying, while at moderate to high temperatures (20 degrees C and 30 degrees C), respiration exhibited rapid declines with decreases in soil moisture. Roots exposed to drying soil also exhibited increased membrane leakage and reduced N. These findings of acclimation of root respiration are important to modelling respiration under different moisture and temperature regimes.     

Context-dependent defence in terrestrial plants: the effects of light and nutrient availability on plant resistance against herbivory

The context‐dependent defence (CDD) hypothesis predicts that defence levels of plant species against herbivory are not fixed but vary with environmental conditions, in a way that is specific for plant species that share evolutionary adaptations to resource conditions exemplified by similar maximum relative growth rates. More specifically, we expected plants from resource‐poor environments to display high defence levels but not when grown under resource‐rich conditions, whereas the reverse – plants from resource‐rich conditions displaying low defence levels but not when grown under resource‐poor conditions – is not necessarily the case. In this study, we used multiple‐choice bioassays in which leaf discs were fed to larvae of Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) as an efficient and effective way of indicating plant defence levels. This generalist herbivore was capable of detecting both inter‐ and intraspecific differences in defence among plant species. The CDD was tested by exploring the effects of various experimental resource conditions (light, nutrients) upon the herbivore preferences and by comparing these preferences with the maximum relative growth rate of plant species. The experimental results provide general support for the CDD hypothesis with respect to nutrient‐level variation but the effects were not related to the origin of the plant species tested. Variation in light conditions did not result in consistent effects upon herbivore preferences. The CDD therefore can be formulated more precisely as: defence levels of plant species vary under different environmental conditions but in a way that is specific for plant species that share evolutionary adaptations to similar nutrient conditions. This more precise CDD hypothesis is a useful addition to existing optimal‐defence theory because of its focus on the possible plastic effects of resource conditions upon plant defence levels. This is relevant when designing experimental plant–herbivore studies.