Effects of soil mechanical impedance on root and shoot growth of Lolium perenne L., Agrostis capillaris and Trifolium repens L.

A method was used for applying a uniform mechanical impedanceto plant roots using sand packed at different bulk densitieswith depth within each growth cylinder. In a growth experiment,replicate cylinders were packed with sand to give the followingmechanical impedances: 0.25 MPa penetration resistance (negligibleimpedance), 1.40 MPa (moderate impedance) and 2.30 MPa (severeimpedance). Seedlings of Lolium perenne L., Trifolium repensL. and Agrostis capillaris were grown for 23 d in each impedancetreatment and effects on both roots and shoots were studied.Severe mechanical impedance affected both root and shoot growthrates for all three species resulting in smaller leaves andshorter roots. For the grasses, the root-to-shoot ratio at harvestwas the same for all the treatments, but a delay in the initiationof both shoots and roots was observed in the severe mechanicalimpedance treatment. The results are discussed in relation tothe possibility that roots penetrating the impeding treatmentsmay have caused signalling that kept shoot and root growth insynchrony. Differences observed in the response to mechanicalimpedance of T. repens compared to the other two species maybe a function of differences in the physiology between mono-and dicotyledonous species. Key words: Mechanical impedance, Lolium perenne, Trifolium repens, roots, shoots     

Membrane Potentials in the Xylem in Roots of Intact Plants

The membrane potential differences (PDs) of root cells of intact,illuminated Trifolium repens L. and Lolium perenne L. have beenmeasured. In T. repens the PDs were the same for all cell typesexcept for the xylem vessels, which were more positive, andfor some cells immediately adjacent to the xylem vessels whichwere 10 mV more negative. The mean PD for all cells was emdash164.6 ± 0.6 mV and the mean for cells adjacent to thexylem vessels with elevated PDs was 178.4 ± 2.4 mV. Whenthe electrode tip was in a xylem vessel a low but stable PD(mean = emdash 89.9 mV) was recorded. The results for L. perennewere similar except that there were no cells with elevated PDsadjacent to the xylem vessels. An inhibitor of ion transport from the root to the shoot, p-fluorophenylalanine(p-FPA), caused a depolarization of 10 mV in the cell PDs butin the xylem vessels the depolarization was 50 mV. The possibility that the elevated PDs of cells adjacent to thexylem vessels are related to the transport of ions into thevessels is discussed.     

The Influence of Volatile Substances from Incubated Litter of Pinus radiata on Seed Germination

Vapour from incubated Pinus radiata D. Don litter caused a depressionin the germination of Lolium perenne L. seeds and responsesvarying from stimulation to depression in germination of Trifoliumrepens L. seeds. These effects could be attributed to ethyleneand carbon dioxide. Experiments using ethylene and carbon dioxidesupported this conclusion. Pinus radiata, litter, Lolium perenne, Trifolium repens, seed germination, ethylene, carbon dioxide     

Comparative analysis of Cd and Zn impacts on root distribution and morphology of Lolium perenne and Trifolium repens: implications for phytostabilization

Backgrounds and aims

The phytostabilization potential of plants is a direct function of their root systems. An experimental design was developed to investigate the impact of Cd and Zn on the root distribution and morphology of Lolium perenne and Trifolium repens.

Methods

Seedlings were transplanted into columns filled with washed quartz and irrigated daily with Cd- or Zn-containing nutrient solutions during 1 month. Root biomass, root length density (RLD) and diameter were subsequently quantified as a function of depth. Pot experiments were also performed to quantify metal, lignin and structural polysaccharides concentrations as well as cell viability.

Results

Lolium perenne accumulated Cd and Zn in the roots whereas T. repens was unable to restrict heavy metal translocation. Cadmium and Zn reduced rooting depth and RLD but induced thick shoot-borne roots in L. perenne. Cd-induced root swelling was related to lignification occurring in the exodermis and parenchyma of central cylinder. Hemicelluloses and lignin did not play a key role in root metal retention. Cadmium slightly reduced mean root cell viability whereas Zn increased this parameter in comparison to Cd.

Conclusions

Even though plant species like Lolium perenne and Trifolium repens may appear suitable for a phytostabilization scheme based on their shoot metal tolerance, exposure to toxic heavy metals drastically impairs their root distribution. This could jeopardize the setting up of phytostabilization trials. The metal-induced alterations of root system properties are clearly metal- and species-specific. At sites polluted with multiple metals, it is therefore recommended to first test their impact on the root system of multiple plant species so as to select the most appropriate species for each site.     

Effects of CO2 enrichment and intraspecific competition on biomass partitioning, nitrogen content and microbial biomass carbon in soil of perennial ryegrass and white clover

Seedlings of perennial ryegrass (Lolium perenne L. cv. Parcour)and white clover (Trifolium repens L. cv. Karina) grown at fivedifferent plant densities were exposed to ambient (390 ppm)and elevated (690 ppm) CO₂ concentrations. After 43 d the effectsof CO₂ enrichment and plant density on growth of shoot and root,nitrogen concentration of tissue, and microbial biomass carbon(C_mic) in soil were determined. CO₂ enrichment of Lolium perenneincreased shoot growth on average by 17% independent of plantdensity, while effects on root biomass ranged between -4% and+ 107% due to an interaction with plant density. Since tilernumber per plant was unaffected by elevated CO₂, the small responseof shoot growth to CO₂ enrichment was atributed to low sinkstrength. A significant correlation between nitrogen concentrationof total plant biomass and root fraction of total plant drymatter, which was not changed by CO₂ enrichment, indicates thatnitrogen status of the plant controls biomass partitioning andthe effect of CO₂ enrichment on root growth. Effects of elevatedCO₂ and plant density on shoot and root growth of Trifoliumrepens were not significantly interacting and mean CO₂-relatedincrease amounted to 29% and 66%, respectively. However, growthenhancement due to elevated CO₂ was strongest when leaf areaindex was lowest. Total amounts of nitrogen in shoots and rootswere bigger at 690 ppm than at 390 ppm CO₂. There was a significantincrease in C_mic in experiments with both species whereas plantdensity had no substantial effect. Key words: CO₂ enrichment, intraspecific competition, biomass partitioning, Lolium perenne, Trifolium repens, grassland     

The effects of elevated CO2 concentrations on the root growth of Lolium perenne and Trifolium repens grown in a FACE* system

Lolium perenne and Trifolium repens were grown in a Free Air CO₂ Enrichment (FACE) system at elevated (600 μimol mol-¹) and ambient (340 μmol mol-¹) carbon dioxide concentrations during a whole growing season. Using a root ingrowth bag technique the extent to which CO₂ enrichment influenced the growth of L, perenne and T. repens roots under two contrasting nutrient regimes was examined. Root ingrowth bags were inserted for a fixed time into the soil in order to trap roots. It was also possible to follow the mortality of roots in bags inserted for different time intervals. Root ingrowth of both L. perenne and T. repens increased under elevated CO₂ conditions. In L. perenne, root ingrowth decreased with increasing nutrient fertilizer level, but for T. repens the root ingrowth was not affected by the nutrient application rate. Besides biomass measurements, root length estimates were made for T, repens. These showed an increase under elevated CO₂ concentrations. Root decomposition appeared to decrease under elevated CO₂ concentrations. A possible explanation for this effect is the observed changes in tissue composition, such as the increase in the carbon: nitrogen ratio in roots of L. perenne at elevated CO₂ concentrations.     

A Comparison of Dry Matter Partitioning in Seminal and Adventitious Plants of Trifolium repens L

Trifolium repens has two types of root, one derived from theseed and the adventitious roots derived from the stolon nodes.It has been suggested that these two systems have differentpotentials for supporting growth. This paper presents a comparisonof plants grown on single seedling or adventitious roots anddemonstrates that although the shoot: root ratios for the twotypes differ this may be explained by differing shoot morphologies.Comparison of the lamina: root ratios for the two types of plantproduced no statistically significant differences and it isproposed that the two types of root system do not differ intheir relationship with leaf growth. A mechanism for large diameter‘tap’ root formation is suggested. white clover, Trifolium repens L., adventitious roots, seedling roots, shoot: root ratio     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions I. Carbon Assimilation and Utilization

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and their growth and carbon economyexamined. The swards received a nutrient solution daily, whichcontained either High (220 mg l^–1) or Low (10 mg l^–1)nitrate N. Rates of canopy photosynthesis and respiration, and final drymatter yields were similar in the two treatments although theproportions of grass and clover differed greatly. The Low-Nswards were made up largely of clover. The grass plants in theseswards had high root: shoot ratios and low relative photosyntheticrates – both signs of N deficiency – and were clearlyunable to compete with the vigorously growing Low-N clover plants.These had higher relative growth rates and dry matter yieldsthan their High-N counterparts. In the High-N swards clovercontributed around 50 per cent to the sward dry weight throughoutthe measurement period despite having a smaller proportion ofits dry weight in photosynthetic tissue (laminae) than grassover much of it. The latter was compensated for, initially bya higher specific leaf area than grass, and later by a higherphotosynthetic rate per unit leaf weight. The results are discussedin relation to observed declines in the clover content of swardsafter the addition of nitrogen fertilizer in the field. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, nitrogen, photosynthesis, carbon balance     

Combined effects between temporal heterogeneity of water supply,nutrient level,and population density on biomass of four broadly distributed herbaceous species

Temporal heterogeneity of water supply affects grassland community productivity and it can interact with nutrient level and intraspecific competition. To understand community responses, the responses of individual species to water heterogeneity must be evaluated while considering the interactions of this heterogeneity with nutrient levels and population density. We compared responses of four herbaceous species grown in monocultures to various combinations of water heterogeneity, nutrient level, and population density: two grasses (Cynodon dactylon and Lolium perenne), a forb (Artemisia princeps), and a legume (Trifolium repens). Treatment effects on shoot and root biomass were analyzed. In all four species, shoot biomass was larger under homogeneous than under heterogeneous water supply. Shoot responses of L. perenne tended to be greater at high nutrient levels. Although root biomass was also larger under homogeneous water supply, effects of water heterogeneity on root biomass were not significant in the grasses. Trifolium repens showed marked root responses, particularly at high population density. Although greater shoot and root growth under homogeneous water supply appears to be a general trend among herbaceous species, our results suggested differences among species could be found in the degree of response to water heterogeneity and its interactions with nutrient level and intraspecific competition.     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions II. Nitrogen Fixation and Nitrate Uptake

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and harvested at intervals over and88 d growht period. The swards received a nutrient solutiondaily, which was either High (220 mg l^–1) or Low (10 mgl^–1) in nitrate N. The nitrate was labelled with the ¹⁵Nisotope. An acetylene reduction assay was carried out on eachsward just prior to harvest. Rates of acetylene reduction agreed qualitatively with the ^l5Nanalyses but absolute values did not match (assuming a 4:1 C₂H₄:N₂ratio) and errors in the acetylene assay are discussed. In theLow-N swards clover relied almost entirely on symbioticallyfixed N₂, fixing more than ten times as much as the High-N cloverplants. In the Low-N treatment the grass was N-deficient despiteobtaining much more nitrate per unit root dry weight than clover.In the High-N swards, however, clover took up more nitrate perunit root weight than grass. The High-N clover plants also fixedsome N₂ and maintained a higher total-N content than grass throughoutthe period. There was no evidence of transfer of symbioticallyfixed N from the clover to the grass in either treatment. Trifolium repens, Lolium perenne, nitrate, nitrogen fixation, ¹⁵N, acetylene reduction     

The Use of a Closed System Flow-Through Enclosure Apparatus for Studying the Effects of Partial Pressure of Dinitrogen in the Atmosphere on Growth of Trifolium repens L. and Lolium perenne L.

A closed system flow-through enclosure apparatus was constructedand used to enclose mixtures ofLolium perenne L. (perennialryegrass cv. Trani) and nodulatedTrifolium repens L. (whiteclover cv. Blanca) growing in soil in pots. There were no significantdifferences between the shoot growth, in terms of dry matteraccumulation and nitrogen content, of mixtures in the systemcompared to that of mixtures grown in a standard growth cabinet.This demonstrated that stable ambient conditions could be achievedby the closed system and its control circuits and that therewere no apparent side-effects of the recirculatory gases.Reducingthe partial pressure of dinitrogen in the atmosphere affectedwhite clover, but not perennial ryegrass. A fairly rapid effectwas observed 4 h after reduction in dinitrogen partial pressureas some of the clover leaves folded downwards along the petiole.These same effects were observed at two different partial pressuresof dinitrogen (22% and 39%) and with two different replacementgases (argon and helium). In the longer term (11 to 16 d) drymatter accumulation and nitrogen content of the clover shootswere significantly reduced. These effects of reduced partialpressure of dinitrogen were observed in both nodulated (NOD⁺)and nitrate-dependent (NOD^–) clover. Possible reasonsfor these effects were discussed with particular reference toimpurities in the gases used, stomatal responses and plant waterrelations. It was concluded that the closed system flow-throughapparatus provides a useful tool for studying whole plant-soilsystems and, in particular, the cycling of nitrogen. However,the use of a replacement gas to reduce the cost of labelleddinitrogen was obviously not a viable proposition. Key words: Dinitrogen, flow-through system, Lolium perenne, partial pressure, Trifolium repens     

The Effects of Rootzone Salinity and Hypoxia on Shoot and Root Growth in Trifolium Species

The effect of concurrent salinity (0-60 mM NaCl) and rootzonehypoxia (flooding for up to 15 d) on shoot and root growth andshoot ion concentrations of six species of Trifolium (T. subterraneumL., T. fragiferum L., T. michelianum Savi., T. isthmocarpumBot., T. purpureum Lois., and T. repens L.), was studied intwo greenhouse experiments. There was a significant salinityx flooding effect for shoot yield but no significant salinityx flooding x species interaction although individual speciesdiffered significantly (P < 0·001) in their growthresponse to the saline or flooded conditions separately. Concentrationsof Na and Cl in the shoots of all species increased with increasingperiods of saline flooding and there was a significant salinityx flooding interaction. Sodium and Cl concentrations were significantlyhigher (P < 0·001) in T. purpureum, the species inwhich shoot growth was most depressed by saline flooding, thanother species. In T. michelianum, T. fragiferum and T. repens,fresh and dry weight of roots increased with flooding underboth saline and non-saline conditions while in T. subterraneumroot growth decreased. A significant proportion of the increasedroot growth in the first three species occurred as new adventitiousroots. These roots had higher percentages of internal gas spaceswithin the root tissue even in the presence of salinity comparedwith roots from non-flooded conditions. There were also significantlymore gas spaces in the total root tissue in T. fragiferum andT. repens under saline-flooding than in roots of T. subterraneum.Electron micrographs of the root cross sections illustratedthe presence of these gas spaces or aerenchyma. Trifolium fragiferum, T. repens and T. michelianum are morelikely to be suited to growth in soils prone to high salinityand to flooding than are T. subterraneum, T. purpureum and T.ishmocarpum.Copyright 1993, 1999 Academic Press Trifolium subterraneum, Trifolium fragiferum, Trifolium michelianum, Trifolium isthmocarpum, Trifolium purpureum, Trifolium repens, salinity, flooding, hypoxia, adventitious roots, aerenchyma, subterranean clover, white clover, strawberry clover, purple clover, balansa clover     

Effects of Edaphic Factors on the Soluble Carbohydrate Content of Roots of Lolium perenneL. and Trifolium repens L.

In a 3³x2² factorial experiment with Lolium perenne L. and Trifoliumrepens L. grown at three soil moisture tensions, three levelsof nitrogen and three levels of phosphorus, there was a widerange of percentage soluble carbohydrate (TSC) in the roots.The significant positive correlations between percentage ofTSC and root weight suggest that root growth-rate was proportionalto the concentration of TSC in the roots. In other experiments with Lolium perenne grown at five constantsoil temperatures and factorial combinations with three soilmoisture treatments with and without manure, there was declinein the percentage of TSC with increasing temperature. This impliedthat the respiration sink strength in the roots did not controlthe partitioning of photosynthate to the roots. No firm conclusions can be drawn from this work, but it is inferredthat there is a dynamic balancing mechanism in the foliage whichcontrols the partitioning of photosynthate between foliage androot growth. This mechanism appears to divert carbohydrate toroots in inverse proportion to root activity.     

Gustatory discrimination between plants by larvae of the scarabaeid, Sericesthis geminata (Coleoptera)

Third-instar larvae of the scarabaeid Sericesthis geminata, given choices between soil containing no root material and soil containing root material of one of the pasture plants Lolium perenne, Phalaris tuberosa or Trifolium repens, accumulated in the soil containing the root material. In addition the larvae responded to differences between plant species and within a plant species.Alcohol and ethyl ether soluble components of plant root extracts generally stimulated feeding. Tests with alcohol, ethyl ether and non-lipid free ethyl ether extracts of roots of the above three plant species and of Dactylis glomerata showed that all except the ethyl ether extract of T. repens were stimulating. There were no other significant interspecific differences between extracts when plants were grown during the summer under the same conditions.

---

Zusammenfassung Die Reaktion von Larven des 3. Stadiums des Scarabaeiden Sericesthis geminata auf die Wurzeln von vier Weidegrasarten wurde in einer Reihe von Wahlexperimenten untersucht. Die Larven sammelten sich vorzugsweise in Böden an, welche wachsende oder lyophilisierte Wurzeln von Lolium perenne, Phalaris tuberosa oder Trifolium repens enthielten, im Gegensatz zu Böden ohne jegliches Wurzelmaterial. Wurde unter gleichen Wachstumsbedingungen eine Auswahl von Wurzelmaterial geboten, so zeigten die Larven eine Vorliebe für L. perenne bwz. T. repens. Unterschiede im Alter und den Wachstumsbedingungen der Pflanzen beeinflußten die Wahl einer bestimmten Art. Äthanol-, Äthyläther- und von Nichtlipoiden befreite Äthylätherextrakte der drei Graswurzeln, die im Treibhaus in Töpfen gezogen wurden, stimulierten die Nahrungsaufnahme der Larven. Äthylätherextrakte von T. repens hatten keinen bedeutenden Einfluß auf die Nahrungsaufnahme, obwohl der von Nichtlipoiden befreite Alkohol- und Äthylätherextrakt die Nahrungsaufnahme erhöhte. Ätherextrakt von P. tuberosa aus dem Freiland wirkte erheblich abschreckend auf die Nahrungsaufnahme der Larven.

     

The role of local environment and geographical distance in determining community composition of arbuscular mycorrhizal fungi at the landscape scale

Arbuscular fungi have a major role in directing the functioning of terrestrial ecosystems yet little is known about their biogeographical distribution. The Baas-Becking hypothesis (‘everything is everywhere, but, the environment selects'') was tested by investigating the distribution of arbuscular mycorrhizal fungi (AMF) at the landscape scale and the influence of environmental factors and geographical distance in determining community composition. AMF communities in Trifolium repens and Lolium perenne roots were assessed in 40 geographically dispersed sites in Ireland representing different land uses and soil types. Field sampling and laboratory bioassays were used, with AMF communities characterised using 18S rRNA terminal-restriction fragment length polymorphism. Landscape-scale distribution of AMF was driven by the local environment. AMF community composition was influenced by abiotic variables (pH, rainfall and soil type), but not land use or geographical distance. Trifolium repens and L. perenne supported contrasting communities of AMF, and the communities colonising each plant species were consistent across pasture habitats and over distance. Furthermore, L. perenne AMF communities grouped by soil type within pasture habitats. This is the largest and most comprehensive study that has investigated the landscape-scale distribution of AMF. Our findings support the Baas-Becking hypothesis at the landscape scale and demonstrate the strong influence the local environment has on determining AMF community composition.     

Uptake of Phosphate by White Clover: I. EVIDENCE FOR AN ELECTROGENIC PHOSPHATE PUMP

Uptake of phosphate from flowing solution and cell membraneelectrical potential differences (PD) have been followed simultaneouslyin the roots of Trifolium repens L. Intact plants were usedand it was found that uptake and PD were strongly influencedby the shoot. They were reduced by excision of the root, defoliationof the plant, and lowering the light intensity at the leaf surface.2, 4-Dinitrophenol caused the PD to decline by approximately40 mV. A close correlation between phosphate uptake and PD wasobtained over a wide range of conditions and removal of phosphatefrom the solution resulted in a decline in PD of about 35 mV.It was concluded that there is an electrogenic phosphate pumpin T. repens roots which is closely dependent on the carbohydratesupply from the shoot.     

Evidence For Root Contraction In White Clover (Trifolium repens L.)

White clover (Trifolium repens L.) stolons become buried inthe field. It was observed that this also occurred in the greenhousewhere the accepted mechanisms of burial, treading by livestockand earthworm casting, did not occur. It was also observed thatthe crown of seedling T. repens plants become closely appressedto the soil. Experiments showed that, regardless of varietyof T. repens or depth of planting, all seedling hypocotyls firstlift the cotyledons clear of the soil, then ‘ contract’towards the soil until the cotyledons are in contact with orbelow the soil surface. Auxanometers were used to measure therate and extent of this contraction and were also attached tostolon nodes in experiments which showed that stolons move downwardsrelative to the soil surface and that the speed and extent ofthis duration varied with soil type. A further experiment showedthat only rooted nodes show this behaviour. The force exertedby the contraction of nodal roots was estimated experimentallyas 0.21 N g^-1fresh root. A mechanism for the root contraction,based on examination of root anatomy of seedling tap-roots andnodal roots, is suggested. These experiments provide evidencefor root contraction in T. repens which may lead to stolon burial.The importance of this to T. repens as a pasture species andas a means of further improving T. repens varieties is discussed.Copyright 1999 Annals of Botany Company White clover, Trifolium repens, L., stolon, seedling, burial, root, nodes, nodal, force, contractile, soil resistance, pasture, phloem, fibres.     

Degree of plant root digestion by the larva of the beetle, Costelytra zealandica

The degree of plant root digestion by grass grubs (Costelytra zealandica) has been evaluated by scanning electron microscope examination of plant residues in faeces. Faecal pellets were examined from grubs fed on roots of pure plant species (perennial ryegrass, Lolium perenne, and white clover, Trifolium repens) under soil-free conditions, as well as from naturally occurring grubs on mixed pasture. There was considerable variation in the degree of digestion observed with different plant species. Clover root was more extensively digested with formation of partially digested xylem vessels, whereas ryegrass root appeared to undergo very little digestive breakdown. These results indicate that plant structural carbohydrates undergo limited digestion by the grass grub. Thus grass grubs may require a high turnover of dietary root material, which would explain the highly destructive effect of the larvae on pasture plants.     

Photosynthesis by White Clover Leaves in Mixed Clover/Ryegrass Swards

Measurements of rates of net photosynthesis were made on singleBlanca white clover leaves on plants taken from a field-grown,mixed clover/perennial ryegrass sward during two regrowth periods. Net photosynthesis fell by 20 per cent in the first measurementperiod as leaf area index increased and the grass componentof the crop flowered, but did not change significantly in thesecond measurement period during which the grass remained vegetative. Leaves which had been artificially protected from shading inthe sward did not have significantly different photosyntheticcapacities from leaves in the undisturbed sward, even in thefirst measurement period. As leaf area index and sward height increased, successive cloverpetioles were longer, keeping the newly expanded leaves nearthe top of the sward where they received full light. It is suggestedthat it is this which allows successive clover leaves, unlikethose of vegetative grasses, to attain a high photosyntheticcapacity throughout a growth period. Trifolium repens, Lolium perenne, Photosynthetic capacity, shading, growth     

Effects of earthworms and organic litter distribution on plant performance and aphid reproduction

Human management practices and large detritivores such as earthworms incorporate plant litter into the soil, thereby forming a heterogeneous soil environment from which plant roots extract nutrients. In a greenhouse experiment we investigated effects of earthworms and spatial distribution of ¹⁵N-labelled grass litter on plants of different functional groups [Lolium perenne (grass), Plantago lanceolata (forb), Trifolium repens (legume)]. Earthworms enhanced shoot and root growth in L. perenne and P. lanceolata and N uptake from organic litter and soil in all plant species. Litter concentrated in a patch (compared with litter mixed homogeneously into the soil) increased shoot biomass and ¹⁵N uptake from the litter in L. perenne and enhanced root proliferation in P. lanceolata when earthworms were present. Growth of clover (T. repens) was rather independent of the presence of earthworms and organic litter distribution: nevertheless, clover took up more nitrogen in the presence of earthworms and exploited more ¹⁵N from the added litter than the other plant species. The magnitude of the effects of earthworms and organic litter distribution differed between the plant species, indicating different responses of plants with contrasting root morphology. Aphid (Myzus persicae) reproduction was reduced on P. lanceolata in the presence of earthworms. We suggest that earthworm activity may indirectly alter plant chemistry and hence defence mechanisms against herbivores.