Root herbivore identity matters in plant-mediated interactions between root and shoot herbivores

Plants are simultaneously attacked by a multitude of herbivores that affect plant responses and plant-mediated interactions in a variety of ways. So far, studies on indirect interactions between below- and aboveground herbivores have almost exclusively focused on interactions between only one root and one shoot herbivore species at the same time. Since these studies show a variety of outcomes, we test the hypothesis that root herbivore identity matters in below-/aboveground interactions. We studied the combined effects root-feeding nematodes (Pratylenchus penetrans) and wireworms (Agriotes lineatus larvae) on Plantago lanceolata and on the performance of aboveground phloem-feeding aphids (Myzus persicae) and chewing caterpillars (Chrysodeixis chalcites larvae). Since root herbivores may also affect resource availability and the microbial community in the rhizosphere, we examined resource utilization by soil microorganisms using BIOLOG EcoPlates™.

Wireworms decreased root biomass by 13%, but led to compensatory shoot growth. Nematodes and the aboveground herbivores did not affect the biomass of Plantago lanceolata. Feeding by C. chalcites larvae enhanced the concentration of aucubin in leaves, which might explain the high mortality of the caterpillars. Aphids and the belowground herbivores did not change iridoid glycoside levels in the leaves. However, the number of aphid offspring was reduced by 44% when nematodes had been added to the soil, whereas wireworms had no effect. We observed higher utilization of BIOLOG carbon sources by the soil microorganisms only in the presence of Pratylenchus penetrans. Our results suggest that the outcome of below–aboveground interactions highly depends on herbivore identity.     

Synergistic effects of antagonistic fungi and a plant growth promoting rhizobacterium, an arbuscular mycorrhizal fungus, or composted cow manure on populations of Meloidogyne incognita and growth of tomato

Effects of four antagonistic fungi (Paecilomyces lilacinus, Pochonia chlamydosporia, Trichoderma harzianum and Gliocladium virens) alone and together with a plant growth promoting rhizobacterium Pseudomonas putida, an arbuscular mycorrhizal fungus Glomus intraradices or with composted cow manure (CCM) were assessed on the growth of tomato and on the reproduction of Meloidogyne incognita in glasshouse experiments. Application of all antagonistic fungi (except G. virens), P. putida, G. intraradices or CCM caused a significant increase in the growth of plants without nematodes. However, use of either of these fungi, P. putida, G. intraradices and CCM against plants with nematodes caused a significant increase in tomato growth. Paecilomyces lilacinus caused a 42% increase in the growth of nematode-inoculated plants followed by P. chlamydosporia (36%), T. harzianum (18%) and G. virens (15%). CCM caused about 57% increase in the growth of nematode-inoculated plants followed by P. putida (37%) and G. intraradices (31%). Maximum increase (71%) in the growth of nematode-inoculated plants was observed when CCM was used with P. lilacinus. Moreover, P. lilacinus caused a high reduction (55%) in galling and nematode multiplication, while G. virens the least (25%). Use of P. putida also caused a 39% reduction in galling and nematode multiplication followed by CCM (34%) and G. intraradices (32%). Combined use of CCM with P. lilacinus caused maximum reduction (79%) in galling and nematode multiplication. Re-isolation of antagonistic fungi from nematodes revealed that P. lilacinus parasitised more females and eggs than other antagonistic fungi. Root colonisation by P. putida was increased with P. lilacinus, while colonisation by G. intraradices was reduced in the presence of antagonistic fungi.     

不同土壤线虫功能团对水稻生长及地上部植食者的影响

探明土壤生物多营养级相互作用是了解生态功能调控机制的核心。本研究通过调控土壤线虫的典型功能团的完全交互设计(食微线虫有无、植食线虫有无、捕杂食线虫有无)探索了线虫功能团对水稻(Oryza sativa)生长及褐飞虱(Nilaparvata lugens)数量的影响。结果表明, 与不接种线虫相比, 植食线虫显著增加水稻根系生物量(P < 0.05), 显著增加其茎叶总酚含量(P < 0.05); 单独的食微线虫增加了褐飞虱数量(P < 0.05), 但显著降低水稻根系生物量(P < 0.05); 捕杂食线虫促进水稻茎叶生长, 降低了褐飞虱数量; 当食微、植食和捕杂食线虫同时存在时, 植物茎叶及根系总酚含量均处于较高水平, 暗示其抗虫潜力更强。总之, 处于较高营养级的捕杂食线虫能够通过调控植食和食微线虫的数量, 提高植物的防御能力, 暗示土壤生物调控措施在植物地上部病原物防控方面有重要的前景。     

Inter- and intra-specific cuticle variation between amphimictic and parthenogenetic species of root-knot nematode (Meloidogyne spp.) as revealed by a bacterial parasite (Pasteuria penetrans)

Specific host–parasite interactions exist between species and strains of plant parasitic root-knot nematodes and the Gram-positive bacterial hyperparasite Pasteuria penetrans. This bacterium produces endospores that adhere to the cuticle of migrating juveniles, germinate and colonise the developing female within roots. Endospore attachment of P. penetrans populations to second-stage juveniles of the root-knot nematode species Meloidogyne incognita and Meloidogyne hapla showed there were interactive differences between bacterial populations and nematode species. Infected females of M. incognita produced a few progeny which were used to establish two nematode lines from single infective juveniles encumbered with either three or 26 endospores. Single juvenile descent lines of each nematode species were produced to test whether cuticle variation was greater within M. hapla lines that reproduce by facultative meiotic parthenogenesis than within lines of M. incognita, which reproduces by obligate parthenogenesis. Assays revealed variability between broods of individual females derived from single second-stage juvenile descent lines of both M. incognita and M. hapla suggesting that progeny derived from a single individual can differ in spore adhesion in both sexual and asexual nematode species. These results suggest that special mechanisms that produced these functional differences in the cuticle surface may have evolved in both sexually and asexually reproducing nematodes as a strategy to circumvent infection by this specialised hyperparasite.     

Above- and belowground insect herbivores differentially affect soil nematode communities in species-rich plant communities

Interactions between above‐ and belowground invertebrate herbivores alter plant diversity, however, little is known on how these effects may influence higher trophic level organisms belowground. Here we explore whether above‐ and belowground invertebrate herbivores which alter plant community diversity and biomass, in turn affect soil nematode communities. We test the hypotheses that insect herbivores 1) alter soil nematode diversity, 2) stimulate bacterial‐feeding and 3) reduce plant‐feeding nematode abundances. In a full factorial outdoor mesocosm experiment we introduced grasshoppers (aboveground herbivores), wireworms (belowground herbivores) and a diverse soil nematode community to species‐rich model plant communities. After two years, insect herbivore effects on nematode diversity and on abundance of herbivorous, bacterivorous, fungivorous and omni‐carnivorous nematodes were evaluated in relation to plant community composition. Wireworms did not affect nematode diversity despite enhanced plant diversity, while grasshoppers, which did not affect plant diversity, reduced nematode diversity. Although grasshoppers and wireworms caused contrasting shifts in plant species dominance, they did not affect abundances of decomposer nematodes at any trophic level. Primary consumer nematodes were, however, strongly promoted by wireworms, while community root biomass was not altered by the insect herbivores. Overall, interaction effects of wireworms and grasshoppers on the soil nematodes were not observed, and we found no support for bottom‐up control of the nematodes. However, our results show that above‐ and belowground insect herbivores may facilitate root‐feeding rather than decomposer nematodes and that this facilitation appears to be driven by shifts in plant species composition. Moreover, the addition of nematodes strongly suppressed shoot biomass of several forb species and reduced grasshopper abundance. Thus, our results suggest that nematode feedback effects on plant community composition, due to plant and herbivore parasitism, may strongly depend on the presence of insect herbivores.     

Interactions between root and shoot herbivores of Ammophila arenaria in the laboratory do not translate into correlated abundances in the field

Over the past decades a growing body of literature has presented proof of the possible interactions between foliar and root herbivores. These effects can be positive, negative or neutral in either direction, depending on the species and the involved mechanism. Most of these studies however concern experiments under controlled conditions. Whether these interactions affect the distribution of herbivores under natural conditions still largely remains an open question. This study examined interactions between root feeding nematodes and shoot feeding aphids on Ammophila arenaria in the laboratory. We subsequently addressed the question whether expectations from this experiment are reflected in correlations between plant related variables and the abundance of both herbivores in the field. We demonstrated that nematodes and aphids can negatively affect each other in a controlled microcosm. In the field however no significant correlations between nematode and aphid abundances could be detected. There, shorter plants with a more vital leaf set and a higher root density supported the highest numbers of aphids. Plants with a lower root density and higher root vitality held more migratory endoparasitic nematodes, while more nematode cysts were found among roots with a low vitality. A certain plant property can furthermore affect above‐ and belowground herbivores in the opposite direction, such as root density in this case. This study suggests that effects of root herbivores on foliar herbivores or vice versa seem to be blurred in a field situation where other variables related to plant vitality and water content structure the herbivore populations. Therefore, caution should be used in generalising the prevalence of these interactions between the above‐ and belowground fauna, based solely on laboratory experiments.     

Mechanism of control of root-feeding nematodes by mycorrhizal fungi in the dune grass Ammophila arenaria

Root-feeding herbivores can affect plant performance and the composition of natural plant communities, but there is little information about the mechanisms that control root herbivores in natural systems. This study explores the interactions between the pioneer dune grass Ammophila arenaria, arbuscular mycorrhizal fungi (AMF) and the root-feeding nematode Pratylenchus penetrans. Our objectives were to determine whether AMF can suppress nematode infection and reproduction and to explore the mechanisms of nematode control by AMF. A sequential inoculation experiment and a split-root experiment were designed to analyse the importance of plant tolerance and resistance and of direct competition between AMF and P. penetrans for the root herbivore and the plant. Root infection and multiplication of P. penetrans were significantly reduced by the native inoculum of AMF. Plant preinoculation with AMF further decreased nematode colonization and reproduction. Nematode suppression by AMF did not occur through a systemic plant response but through local mechanisms. Our results suggest that AMF are crucial for the control of root-feeding nematodes in natural systems and illustrate that locally operating mechanisms are involved in this process.     

Induced resistance by the mutualistic endophyte, Fusarium oxysporum strain 162, toward Meloidogyne incognita on tomato

The non-pathogenic endophytic fungus, Fusarium oxysporum strain 162, originally isolated from the endorhiza of tomato roots, reduces damage caused by Meloidogyne incognita, by inhibiting juvenile penetration of and development in the root. However, little is known about the mode of action of this endophyte fungus against the nematode. This study aimed at investigating how the endophyte affects nematode motility and survival and if induced resistance plays a role in the relationship. In a previous study, F. oxysporum strain 162 decreased nematode penetration of tomato up to 60%. In experiments using a split-root chamber to test for induced resistance, nematode penetration, number of galls, and number of egg masses were investigated 2 and 5 weeks after nematode inoculation. Split-root plants treated with F. oxysporum strain 162 showed 26-45% less nematode penetration, 21-36% less galls and a 22-26% reduction in the number of egg masses in the roots not directly inoculated with the fungus when compared to untreated control plants in repeated tests. In conclusion, inoculation of tomato plants with the non-pathogenic fungal endophyte F. oxysporum strain 162 resulted in a signficant reduction of nematode infection, which was in part due to induced resistance in the first 2-3 weeks after fungal inoculation.     

Interactions of Paecilomyces lilacinus strain 251 with the mycorrhizal fungus Glomus intraradices: Implications for Meloidogyne incognita control on tomato

The interactions of Paecilomyces lilacinus strain 251 with the arbuscular mycorrhizal fungus Glomus intraradices and their significance for the control of Meloidogyne incognita on tomato were investigated in greenhouse experiments. Application of P. lilacinus had no effect on the frequency and intensity of tomato root colonization by G. intraradices. Likewise, the decline of the nematophagous fungus densities after single application in soil was not affected by the presence of the mycorrhizal fungus. Single application of P. lilacinus, as pre-planting soil treatment, resulted in significant reduction of nematode damage. In contrast, mycorrhizal inoculation did not provide sufficient biocontrol. Combined application of the two agents did not enhance root protection compared to single treatments. Double treatment of mycorrhized seedlings with P. lilacinus, as seedling drench and pre-planting soil treatment, 4 and 1 week before transplanting, respectively, resulted in the highest reduction of the nematode damage. These results indicate the potential of the commercial P. lilacinus strain 251 and mycorrhiza for integration in nematode control strategies.     

Effects of competition on an endangered limpet Patella ferruginea (Gastropoda: Patellidae): Implications for conservation

Spatial and trophic competitive interactions were analysed for the seldom-studied limpet Patella ferruginea (which is considered as an “extinction risk”). Spatial patterns of distribution of P. ferruginea were studied using quadrats of 1 × 1 m² at several transects along the coastline, whereas trophic interactions were studied through manipulations of densities of P. ferruginea and Patella caerulea in experimental plots located at the mid-shore. Physical parameters were measured inside each plot to determine the microenvironmental differences between fenced and unfenced plots. Fenced plots of 50 × 50 cm² were used with six different treatments. Mortality was recorded every 15 days and length was measured monthly during the 3.5 months of the experiment. At the beginning and end of the experiment, lengths and weights of limpets were recorded and chlorophyll concentration was measured by spectrophotometric analysis of rock surface. Both species showed increased mortality and reduced size and weight in treatments with greater densities. High intraspecific competition was detected in P. caerulea compared with P. ferruginea whereas asymmetrical interspecific competition was observed. The results suggest that P. caerulea is the superior competitor, possibly due to differences in radular morphology and mobility between the two species. Microalgal food assessment showed that it is probably a limiting resource, especially for P. caerulea, although this species has a lower biomass than P. ferruginea. The high intraspecific competition observed in P. caerulea could be responsible for the coexistence of both species. The present study may be useful in helping to conserve P. ferruginea, which is presently under serious threat of extinction, because the results show that experimental fences are a viable means of concentrating breeding individuals.     

The Influence of Trichoderma harzianum on the Root-knot Fusarium Wilt Complex in Cotton

Wilt-susceptible cultivar ''Rowden'' cotton was inoculated wilh Meloidogyne incognita (N), Trichoderma harzianum (T), and Fusarium oxysporum f. sp. vasinfectum (F) alone and in all combinations in various time sequences. Plants inoculated with F alone or in combination with T did not develop wilt, Simultaneous inoculation of 7-day-old seedlings with all three organisms (NTF) produced earliest wilt. However, plants receiving nematodes at 7 days and Fusarium and Trichoderma at 2 or 4 weeks later (N-T-F, N-TF) developed the greatest wilt between 49-84 days after initial nematode inoculation. During the same period, Fusarium added 4 weeks after initial nematode inoculation (N-F) and Fusarium added 4 weeks after initial simultaneous inoculation of nematode and Trichoderma (NT-F) produced the least wilt. The addition of Fusarium inhibited nematode reproduction. Simultaneous inoculation with nematodes and Trichoderma (NT-) resulted in the greatest root gall development, whereas nematodes alone produced the greatest number of larvae. In comparison with noninoculated controls (CK), treatments involving all three organisms inhibited plant growth, plants inoculated with the nematode alone (N-) or with nematodes and Trichoderma (NT-) simultaneously had greatest root weight. Any treatment involving the nematode resulted in fewer bolls per plant and greater necrosis on roots than the noninoculated checks.     

Effects of waterlogging and increased soil nutrients on growth and reproduction of Polygonum hydropiper in the hydro-fluctuation belt of the Three Gorges Reservoir Region

水淹和土壤养分是影响三峡库区消落带植物生长的主要环境因子。消落带不同高程的植物长期经历不同的淹水强度和土壤养分条件。该研究假设同一物种来自于消落带不同高程的植株可能产生性状分化, 从而对根部淹水和土壤养分变化具有不同的生长和繁殖响应策略。为了验证以上假设, 选取在三峡库区消落带高低高程均广泛分布的物种水蓼(Polygonum hydropiper)为研究对象, 采集自然种群的种子。在温室同质园条件下, 研究了根部水淹和土壤养分提升对高低高程水蓼植株生长和繁殖特性的影响。研究结果表明根部水淹显著或趋于显著降低了水蓼植株功能叶的叶长、叶宽、总分枝数、叶生物量、花生物量和总生物量; 低养分处理显著或趋于显著降低了水蓼植株的总节数、总分枝数、根生物量、花生物量和总生物量, 表明根部水淹和低土壤养分对水蓼的生长和繁殖能力具有抑制作用。同时, 根部水淹和土壤养分的交互作用显著影响植株的根生物量, 表明根部水淹条件下高土壤养分更有利于植株根生物量的积累。高高程植株的根生物量和叶生物量显著或趋于显著高于低高程植株, 而低高程植株的始花时间早于高高程植株, 且繁殖分配也显著高于高高程植株, 表明高低高程水蓼植株对资源的分配策略不同。该研究结果表明水蓼的生长和繁殖特性受根部水淹和土壤养分共同限制, 但对根部水淹条件下高土壤养分生境具有较好的适应性; 同时, 低高程植株可以通过调整其生长和繁殖特性以提高对所处生境胁迫的适应性。     

Effects of glyphosate on interspecific competition between Solidago canadensis and Imperata cylindrica

Aims The extensive use of herbicide to control invasive plants would change the relationship between alien and neighboring plants. In order to provide data for rational use of herbicide and a theoretical reference for further studies on the ecological effects of glyphosate, we explored the variation of the relationship between an invasive plant Solidago canadensis and a native plant Imperata cylindrica when they were sprayed glyphosate.
Methods A replacement series experiment was conducted from June to August 2016 in Wetland Ecosystem Research Station of Hangzhou Bay, State Forestry Administration, to examine the effects of glyphosate at seven concentration levels (0, 0.3, 0.6, 0.9, 1.2, 1.5, 1.8 mL·L^-1) on the growth and interspecific competition of S. canadensis and I. cylindrica.
Important findings (1) Glyphosate treatment significantly inhibited the growth of S. canadensis and I. cylindrica (p < 0.05). During the test, cumulative growth of height and leaf number of S. canadensis were apparently reduced with the increase of glyphosate concentration, but the leaf number of S. canadensis treated with 0.3- 1.5 mL·L^-1 glyphosate was re-growing with time, while the one treated with 1.8 mL·L^-1 was mostly dead. The withering rate of tiller and green leaf of I. cylindrica also significantly increased with the increase of glyphosate concentration, and the growth indices of this plant treated with 0.3-0.6 mL·L^-1 were also re-growing with time. (2) Glyphosate treatment significantly affected interspecific competition (p < 0.05), which diminished as the glyphosate concentration increased. (3) Interspecific competition has significant influence on the biomass allocation of S. canadensis (p < 0.05). When facing competition, S. canadensis would allocate more organic matter to root and thus increase the ratio of root to shoot. Competition only inhibited the tiller number and total biomass of I. cylindrica, but insignificantly affected its ratio of root to shoot. (4) The interaction between glyphosate treatment and S. canadensis-I. cylindrica interspecific relationship also significantly influenced the biomass of S. canadensis and I. cylindrica (p < 0.05), but insignificantly affected the root/shoot ratio of two plants. Different plants have different tolerance to glyphosate stress. Compared with native plant I. cylindrica, S. canadensis has stronger tolerance to glyphosate. Low-concentration glyphosate could decrease the competitive intensity between S. canadensis and I. cylindrica, which may disturb the structure and dynamics of plant communities.     

Differing interactions between an introduced beetle and a resident root nematode mediated by an invasive plant and its native congener

Interactions between above- and below-ground herbivores play an important role in shaping plant competition and invasion, while the effects of non-native species invasions on above- and below-ground interactions remain unexplored. In this study, we report the interactions between an above-ground introduced beetle and a resident root nematode hosted by an invasive plant or its native congener with a laboratory bioassay and a greenhouse experiment in Wuhan, China. Nematode infections decreased beetle food conversion rates and larval biomass on the native plant, and increased beetle food conversion rates with no detectable impact on the larval biomass on the invasive plant. Beetle defoliation decreased nematode egg production on both the native and invasive plants. The interactions of the introduced beetle and the nematode were different by the invasive and native plants, which suggests that invasive plants and their introduced herbivores have the potential to alter above- and below-ground interactions and affect associated community members, which may in turn affect invasion processes and the safety of classical biocontrol practices.     

Toward a mechanistic understanding of competition in vascular-feeding herbivores: an empirical test of the sink competition hypothesis

Recent evidence suggests that competitive interactions among herbivores are mostly indirect and mediated by plant responses to herbivory. Most studies, however, emphasize chewing insects and secondary chemistry, thus ignoring the diverse group of vascular-parasites that may be more likely to compete through induced changes in phytonutrients. Using an aboveground phloem-feeding aphid (Myzus persicae) and a belowground gall-forming nematode (Meloidogyne incognita) on tobacco plants, we assessed the importance of competition via induced host–plant sinks. In a series of experimental trials, nematode root herbivory caused 55 and 72% declines in the growth and fecundity of aphids, respectively. Aphids, on the other hand, did not impact nematode performance. Therefore, we predicted that nematodes out-compete M. persicae by attenuating the magnitude of aphid-induced sinks. Through a combination of invertase enzyme measurements and stable isotope (¹³C and ¹⁵N) enrichment, we found evidence that both herbivores act as mobilizing sinks. Aphids attracted photoassimilates to feeding aggregations on leaves and nematode galls accumulated resources in the roots. Levels of invertase enzymes, for example, were more than fourfold higher in nematode galls than in surrounding root tissue. Yet we found no evidence supporting a sink competition model for aphid–nematode interactions. The strength of aphid-induced leaf sinks was entirely unaffected by nematode presence, and vice versa. Thus, induced host–plant sinks appear to be a common strategy employed by vascular parasites to manipulate the physiology of their host, but multi-sink competition may be limited to herbivores that co-occur on the same tissue type and/or plants under growth-limited abiotic conditions.     

根部水淹和土壤养分提升对三峡库区消落带水蓼生长和繁殖特性的影响

水淹和土壤养分是影响三峡库区消落带植物生长的主要环境因子。消落带不同高程的植物长期经历不同的淹水强度和土壤养分条件。该研究假设同一物种来自于消落带不同高程的植株可能产生性状分化, 从而对根部淹水和土壤养分变化具有不同的生长和繁殖响应策略。为了验证以上假设, 选取在三峡库区消落带高低高程均广泛分布的物种水蓼(Polygonum hydropiper)为研究对象, 采集自然种群的种子。在温室同质园条件下, 研究了根部水淹和土壤养分提升对高低高程水蓼植株生长和繁殖特性的影响。研究结果表明根部水淹显著或趋于显著降低了水蓼植株功能叶的叶长、叶宽、总分枝数、叶生物量、花生物量和总生物量; 低养分处理显著或趋于显著降低了水蓼植株的总节数、总分枝数、根生物量、花生物量和总生物量, 表明根部水淹和低土壤养分对水蓼的生长和繁殖能力具有抑制作用。同时, 根部水淹和土壤养分的交互作用显著影响植株的根生物量, 表明根部水淹条件下高土壤养分更有利于植株根生物量的积累。高高程植株的根生物量和叶生物量显著或趋于显著高于低高程植株, 而低高程植株的始花时间早于高高程植株, 且繁殖分配也显著高于高高程植株, 表明高低高程水蓼植株对资源的分配策略不同。该研究结果表明水蓼的生长和繁殖特性受根部水淹和土壤养分共同限制, 但对根部水淹条件下高土壤养分生境具有较好的适应性; 同时, 低高程植株可以通过调整其生长和繁殖特性以提高对所处生境胁迫的适应性。     

Relationship Between Paratrichadorus sp. Density, and Growth of Wheat in Pots

The effect of a Paratrichodorus sp. (close to P. tunisiensis) on the growth of wheat (Triticum durum Desf.) was investigated in pots containing different nematode densities and maintained in a growth chamber at 20 C for 40 days. The relation between fresh weight of tops and initial nematode density was according to the equation y = m + (1 - m)z^P⁻T. This suggests a tolerance limit of 1.4 nematodes/cm³ of soil under the conditions of the experiment; taking into account the effect of the great nematode mortality, it is estimated to be between 0.15 and 0.35 nematodes/cm³ soil. Models of the growth of the plants and the multiplication of the nematodes (assuming a constant mortality of the nematodes in the absence of roots) which explain the relation between initial and tinal nematodes densities at initial densities greater than 1 nematode/cm³ soil are described in an appendix. Sections of nematode infested roots showed disorganization of root structure clue to abnormal proliferation of lateral roots. Nematode feeding on the root cap and apical meristem caused cessation of root elongation and induced abnormal production of lateral root primordia.     

Effects of native and exotic range‐expanding plant species on taxonomic and functional composition of nematodes in the soil food web

Due to climate warming, many plant species shift ranges towards higher latitudes. Plants can disperse faster than most soil biota, however, little is known about how range‐expanding plants in the new range will establish interactions with the resident soil food web. In this paper we examine how the soil nematode community from the new range responds to range‐expanding plant species compared to related natives. We focused on nematodes, because they are important components in various trophic levels of the soil food web, some feeding on plant roots, others on microbes or on invertebrates. We expected that range expanding plant species have fewer root‐feeding nematodes, as predicted by enemy release hypothesis. We therefore expected that range expanders affect the taxonomic and functional composition of the nematode community, but that these effects would diminish with increasing trophic position of nematodes in the soil food web. We exposed six range expanders (including three intercontinental exotics) and nine related native plant species to soil from the invaded range and show that range expanders on average had fewer root‐feeding nematodes per unit root biomass than related natives. The range expanders showed resistance against rather than tolerance for root‐feeding nematodes from the new range. On the other hand, the overall taxonomic and functional nematode community composition was influenced by plant species rather than by plant origin. The plant identity effects declined with trophic position of nematodes in the soil food web, as plant feeders were influenced more than other feeding guilds. We conclude that range‐expanding plant species can have fewer root‐feeding nematodes per unit root biomass than related natives, but that the taxonomic and functional nematode community composition is determined more by plant identity than by plant origin. Plant species identity effects decreased with trophic position of nematodes in the soil food web.     

Biomass Partitioning in Tomato Plants Infected with Meloidogyne incognita

Tomato plants were inoculated with Meloidogyne incognita at initial populations (Pi) of 0, 1, 10, 50, 100, and 200 (x 1,000) eggs per plant and maintained in a growth chamber for 40 days. Total fresh biomass (roots + shoots) at harvest was unchanged by nematode inoculation with Pi of 1 x 10⁵ eggs or less. Reductions in fresh shoot weight with increasing Pi coincided with increases in root weight. Total fresh biomass declined with Pi above 1 x 10⁵ eggs, whereas total dry biomass declined at Pi above 1 x 10⁴ eggs. The greatest reduction percentages in fresh shoot biomass induced by root-knot nematodes occurred in the stem tissue, followed by the petiole + rachis; the least weight loss occurred in the leaflets. Although biomass varied among shoot tissues, the relationship between biomass of various shoot tissues and Pi was described by quadratic equations. The linear and quadratic coefficients of the equations (stem, petiole + rachis, or leaflets on Pi) did not differ among tissues when calculations were based on standardized values. Meloidogyne incognita-infected plants had thinner leaves (leaf area/leaf weight) than did uninfected plants. Reductions in leaf weight and leaf area with nematode inoculation occurred at nodes 5-15 and 4, 6-14, respectively. Losses in plant height and mass due to nematodes reflected shorter internodes with less plant mass at each node.