Release of allelochemical agents from litter,throughfall, and topsoil in plantations ofEucalyptus globulus Labill in Spain

Natural leachates ofEucalyptus globulus (throughfall, stemflow, and soil percolates) were collected daily during rainy spells in the vegetative period (February–July), and their effects on the germination and radicle growth ofLactuca sativa were measured. Concurrently, the effects ofL. sativa of topsoil and leachates from decaying litter were determined. The results suggest that toxic allelochemicals released byEucalyptus globulus may influence the composition and structure of the understory of the plantation and that this effect is attributable mainly to the decomposition products of decaying litter rather than to aerial leachates. The soil may neutralize or dilute allelopathic agents, at least below the top few cms.     

Inhibition of seedling growth of crop species by recirculating root exudates ofBidens pilosa L.

The allelopathic effects of root exudates ofBidens pilosa L. on seedling growth ofLactuca sativa L.,Phaseolus vulgaris L.,Zea mays L., andSorghum bicolor (L.) Moench were studied using a root exudate recirculating system that allows continuous exposure of crop plants to allelopathic chemicals. This system maintains an undisturbed rhizosphere and eliminates competition and physical contact between the donor and acceptor plants. Comparison of responses to hydrophobic and hydrophilic root exudates is made possible by removal of hydrophobic compounds using XAD-4. Treatments consisted ofB. pilosa, B. pilosa with an Amberlite XAD-4 resin column attached to the donor pot to remove hydrophobic allelochemicals, and a donor pot without weeds.B. pilosa significantly inhibited seedling growth of all crop species tested. The crop species varied in response to the root exudates, withL. sativa being most sensitive. Larger and olderB. pilosa plants caused greater inhibition of seedling growth ofL. sativa andP. vulgaris than did smaller (younger)B. pilosa plants.B. pilosa with XAD-4 caused significantly less inhibition to all crop species, exceptZ. mays, thanB. pilosa without XAD-4, indicating that the hydrophobic exudates played an important role in the allelopathic growth inhibition. Variability in species response toB. pilosa with and without XAD-4 was probably due to differences in sensitivity to hydophobic and hydrophilic allelochemicals.Supported by a grant under USDA Agreement No. 83-CRSR-2-2293. Journal Series No. 2887 of the Hawaii Institute of Tropical Agriculture and Human Resources.     

The Role of Pentacyclic Triterpenoids in the Allelopathic Effects of Alstonia scholaris

Alstonia scholaris is a tropical evergreen tree native to South and Southeast Asia. Alstonia forests frequently lack understory species. However, potential mechanisms—particularly the allelochemicals involved—remain unclear. In the present study, we identified allelochemicals of A. scholaris, and clarified the role of allelopathic substances from A. scholaris in interactions with neighboring plants. We showed that the leaves, litter, and soil from A. scholaris inhibited growth of Bidens pilosa—a weed found growing abundantly near A. scholaris forests. The allelochemicals were identified as pentacyclic triterpenoids, including betulinic acid, oleanolic acid, and ursolic acid by using ¹H and ¹³C-NMR spectroscopy. The half-maximal inhibitory concentration (IC₅₀) for radicle growth of B. pilosa and Lactuca sativa ranged from 78.8 μM to 735.2 μM, and ursolic acid inhibited seed germination of B. pilosa. The triterpenoid concentrations in the leaves, litter, and soil were quantified with liquid chromatography-electrospray ionization/tandem mass spectrometry. Ursolic acid was present in forest soil at a concentration of 3,095 μg/g, i.e., exceeding the IC₅₀. In the field, ursolic acid accumulated abundantly in the soil in A. scholaris forests, and suppressed weed growth during summer and winter. Our results indicate that A. scholaris pentacyclic triterpenoids influence the growth of neighboring weeds by inhibiting seed germination, radicle growth, and functioning of photosystem II.     

Allelopathic potential of Macaranga tanarius (L.) muell.-arg

Macaranga tanarius is widely distributed in the abandoned lowlands of Taiwan where substantial amounts of leaves accumulate on the ground. A unique pattern of weed exclusion underneath trees is often found and thought to result from allelopathic interactions. Density-dependent phytotoxicity analysis of Lactuca sativa L. (lettuce) growing in soil mixed with the powder of M. tanarius leaves showed a significant deviation from the expected yield–density relationship. Lettuce growth was most suppressed in the low seed density experiment suggesting that the phytotoxins produced during leaf decomposition inhibit the growth of lettuce seedlings. Bidens pilosa and Leucaena leucocephala, growing in soil mixed with the leaf powder of M. tanarius were also suppressed. Aqueous leaf extracts were bioassayed against lettuce and B. pilosa, and exhibited a significant suppression in radicle growth. Compounds identified from leaves included nymphaeol-A (1), nymphaeol-B (2), nymphaeol-C (3), quercetin (4), abscisic acid (ABA) (5), blumenol A (6), blumenol B (7), roseoside II (8), tanariflavanone A (9), and tanariflavanone B (10). ABA was the major growth inhibitor. At concentrations of 20 ppm, ABA suppressed lettuce germination, while at 120 ppm it inhibited the growth of Miscanthus floridulus, Chloris barbata, and Bidens pilosa. At 600 ppm, quercetin, blumenol A, and blumenol B, caused 20–25% inhibition of radicle and shoot growth of M. floridulus. The amount of ABA in M. tanarius leaves was approximately 3–5 g g^–1 dry weight, significantly higher than previously reported. We conclude that the pattern of weed exclusion underneath stands of M. tanarius and its invasion into its adjacent grassland vegetation results from allelopathic interactions.     

Allelopathic research of subtropical vegetation in Taiwan

Leucaena leucocephala plantations in Kaoshu, southern Taiwan, exhibit, after several years of growth, a unique pattern of weed exclusion beneathLeucaena canopy. The pattern has been observed in manyLeucaena plantations in Taiwan and is particularly pronounced in the area where a substantial amount ofLeucaena litter has accumulated on the ground. Field data showed that the phenomenon was primarily not due to physical competition involving light, soil moisture, pH, and nutrients. Instead, aqueous extracts ofLeucaena fresh leaves, litter, soil, and seed exudate showed significantly phytotoxic effects on many test species, including rice, lettuce,Acacia confusa, Alnus formosana, Casuarina glauca, Liquidambar formosana, andMimosa pudica. However, the extracts were not toxic to the growth ofLeucaena seedlings. The decomposing leaves ofLeucaena also suppressed the growth of the aforementioned plants grown in pots but did not inhibit that ofLeucaena plants. By means of paper and thin-layer chromatography, UV-visible spectrophotometry, and high-performance liquid chromatography, 10 phytotoxins were identified. They included mimosine, quercetin, and gallic, protocatechuic,p-hydroxybenzoic,p-hydroxyphenylacetic, vanillic, ferulic, caffeic, andp-coumaric acids. The mature leaves ofLeucaena possess about 5% dry weight of mimosine, the amount varying with varieties. The seed germination and radicle growth of lettuce, rice, and rye grass were significantly inhibited by aqueous mimosine solution at a concentration of 20 ppm, while that of the forest species mentioned was suppressed by the mimosine solution at 50 ppm or above. However, the growth ofMiscanthus floridulus andPinus taiwanensis was not suppressed by the mimosine solution at 200 ppm. The seedlings ofAgeratum conzoides died in mimosine solution at 50 ppm within seven days and wilted at 300 ppm within three days. It was concluded that the exclusion of understory plants was evidently due to the allelopathic effect of compounds produced byLeucaena. The allelopathic pattern was clearly shown in the area with a heavy accumulation ofLeucaena leaf litter, which was a result of drought and heavy wind influence.Paper No. 292 of the Scientific Journal Series of the Institute of Botany, Academia Sinica, Taipei, Taiwan. This study was supported in part by a grant to C.H. Chou. Part of this paper was a MS thesis submitted by Y.L. Kuo to the Department of Forestry, National Taiwan University, and presented at the Seminar on Allelochemicals and Pheromones, sponsored by the CCNAA and AIT on June 21–26, 1982.     

Phytotoxicity of cacalol and some derivatives obtained from the roots ofPsacalium decompositum (A. Gray) H. Rob. & Brettell (Asteraceae), matarique or maturin

The effect of cacalol and extracts (water and petroleum ether) derived from the roots ofPsacalium decompositum (Asteraceae) on the germination and radicle growth of two plants.Amaranthus hypochondriacus (Amaranthaceae) andEchinochloa crusgalli (Poaceae), and on the radial growth of four phytopathogenic fungi is described. The activity of two cacalol derivatives (methyl cacalol and cacalol acetate) was also investigated. Germination ofA. hypochondriacus was inhibited by almost all the treatments. The extracts and cacalol produced a significant inhibition of radicle growth ofA. hypochondriacus andE. crusgalli. Cacalol acetate showed a specific inhibition onE. crusgalli, and methyl cacalol significantly inhibited the growth ofA. hypochondriacus. In general, antifungal activity depended upon the target fungi and the concentration of each treatment. Cacalol had also effects on the morphology and coloration of the fungal mycelium. The bioactivity shown by the extracts ofPsacalium decompositum on the tested seeds and fungi is mainly due to their content in cacalol.     

Role of allelopathy as a possible factor associated with the rising dominance ofBunias orientalis L. (Brassicaceae) in some native plant assemblages

Leaf extracts ofBunias orientalis were shown to inhibit seed germination of a variety of cultivar plant species and of species cooccurring withB. orientalis in the field. Root exudate solutions and leaf litter leachates ofB. orientalis were tested for their allelopathic activity using seedling growth assays. Additionally, in comparative seedling growth assays soil cores removed from denseB. orientalis stands were tested bimonthly for elevated allelopathic effects. The impact of root exudates on seedling growth was generally weak and varied between species. Similar results were obtained for the effect ofB. orientalis leaf litter leachates on seedlings grown in sand culture relative to the effect of leaf litter leachates of a plant species mixture. When soil as a growth substrate was used, no consistent differences in seedling growth were obtained between the two litter leachate treatments. In the soil core experiment seedlings grown in soil cores collected from a denseB. orientalis stand unexpectedly showed better performance than seedlings grown in soil cores collected from a nearby mixed plant stand withoutB. orientalis, at least in early spring and late autumn. Predominating nutrient effects are, therefore, assumed to conceal a potentially increased allelopathic effect of soil beneath denseB. orientalis stands. It is concluded that other factors than allelopathy must be investigated to explain the rapid establishment of dense stands of this alien plant species.     

Allelopathic potential and chemical constituents of volatiles from Ageratum conyzoides under stress

The allelopathic potential of Ageratum conyzoides was investigated under different environmental stress conditions, including nutrient deficiency, physical damage, 2,4-D treatment, competition with Bidens pilosa, infection with Erysiphe cichoracearum, and feeding by Aphiids gossypii. The inhibitory effects of A. conyzoides volatiles on peanut (Arachis hypogaea), redroot amaranth (Amaranthus retroflexus), cucumber (Cucumis sativus), and ryegrass (Lolium multiforum) increased when plants were grown under nutrient-deficient conditions or in competition with B. pilosa; however, there was no difference with physical damage or 2,4-D treatment. Phytoinhibitory effects decreased under fungal infection and aphid feeding. Volatiles from A. conyzoides plants infected with E. cichoracearum or exposed to A. gossypii feeding inhibited or killed fungi and insects. Precocenes and their derivatives, monoterpenes, and sesquiterpenes were the major volatile components of A. conyzoides.     

Allelopathy as a factor in the success ofHelianthus mollis Lam.

A method to rapidly screen species suspected of producing allelochemics, using results from simple bioassay tests, is presented. By measuring the osmotic potential ofH. mollis extracts and using mannitol solutions of comparable osmotic potential, the influence of osmotic potential in the bioassay was eliminated. Nested analysis of variance was used to examine the separate influences of (1) extract concentration, (2) source of plants used in extract preparation (edge or center of clones) (3) osmotic potential of the extract, and (4) the differential development of radicles and shoots of species used in the bioassay tests. Bioassay tests for allelopathy showed that extracts made of wholeH. mollis plants significantly inhibited both radicle and shoot development of radish and wheat, but only the radicle of little bluestem. There was a significant increase in the inhibition of radish shoots and wheat radicles at high concentration of the extract, but the radicle of little bluestem was inhibited more at the lower concentration. Extracts prepared from plants collected from the clone center inhibited radish radicle development significantly more than extracts made of plants growing at the clone edge.     

Allelopathic research of subtropical vegetation in Taiwan II. Comparative exclusion of understory byPhyllostachys edulis andCryptomeria japonica

On many hillsides of Taiwan there is a unique pattern of weed exclusion byPhyllostachys edulis (bamboo) andCryptomeria japonica (conifer) in which the density, diversity, and dominance of understory species are very different. Although the physical conditions of light, soil moisture, and soil nutrients strongly favor the growth of understory in a bamboo community, the biomass of its undergrowth is significantly low, indicating that physical competition among the understory species in the bamboo and conifer communities does not cause the observed differences. However, the biochemical inhibition revealed by these two plants appeared to be an important factor. The growth ofPellionia scabra seedlings, transplanted from the study site into greenhouse pots, was evidently suppressed by the aqueous leachate of bamboo leaves but was stimulated by that of conifer leaves. The radicle growth of lettuce, rye grass, and rice plants was also clearly inhibited by the leachate and aqueous extracts of bamboo leaves but not by those of conifer leaves. Six phytotoxins,o-hydroxyphenylacetic,p-hydroxybenzoic,p-coumaric, vanillic, ferulic, and syringic acids were found in the aqueous leachate and extracts of leaves and alcoholic soil extracts ofP. edulis, while the first three compounds were absent in the extracts ofC. japonica. The phytotoxicities of extracts were correlated with the phytotoxins present in both leaves and soils. The understory species might be variously tolerant to the allelopathic compounds produced by the two plants, resulting in a differential selection of species underneath. Therefore, comparative allelopathic effects ofPhyllostachys edulis andCryptomeria japonica may play significant roles in regulating the populations of the understories.Paper No. 253 of the Scientific Journal Series of the Institute of Botany, Academia Sinica, Taipei, Taiwan. This study was supported by the National Science Council of the Republic of China.     

Allelopathic potential of corn pollen

The effects of corn pollen and aqueous leachates of pollen upon the radicle growth ofBidens pilosa, Cassia jalapensis, andRumex crispus are shown. Extractions of pollen with various solvents and methods were carried out so as to assess its active principle and its effect uponC. jalapensis. The preliminary steps to separate and identify the allelopathic compounds of the sonicated and macerated pollen extracted with methylene chloride are described. The strongest inhibitory effect was produced by the hexane fraction. The allelopathic effect of corn pollen upon the growth ofC. jalapensis in several substrates is shown. The possible structure of some of the active fractions is discussed as well as the possibility that the allelopathic potential of pollen might actually occur in nature.     

Evidence for allelopathy by tree-of-heaven (Ailanthus altissima)

Ailanthus altissima (Mill.) Swingle contains one or more phytotoxic compounds in roots and leaves. Activity is higher in roots, where it occurs primarily in the bark. Powdered root bark and leaflets strongly inhibited growth of garden cress (Lepidium sativum L.) when mixed with soil in Petri dishes (ID₅₀ values=0.03 g root bark, 0.6 g leaflet/dish). The toxic material was readily extracted by methanol but not dichloromethane. Pieces of root bark mixed with soil at 2, 1, and 0.5 g/pot reduced cress biomass in the greenhouse, whereas methanol-extracted root bark did not. The inhibitory effect ofAilanthus tissues in soil was short-lived (4 weeks in pots in greenhouse, 3 days in Petri dishes in laboratory). Inhibition by root bark was sometimes superseded by stimulation. FreshAilanthus root segments placed in or on soil reduced growth of nearby cress seedlings. Fine roots were more inhibitory than coarse, and inhibition became more pronounced with increased time of soil exposure to roots. Soil collected nearAilanthus roots in the field supported reduced radicle growth of cress compared to control soil. In contrast, stemflow fromAilanthus trees stimulated cress growth. The results suggest allelopathy caused by toxin exudation from roots may contribute to the aggressiveness and persistence ofAilanthus in certain habitats.     

Clinical and biochemical effects of a combination botanical product (ClearGuard™) for allergy: a pilot randomized double-blind placebo-controlled trial

Background  

Botanical products are frequently used for treatment of nasal allergy. Three of these substances, Cinnamomum zeylanicum, Malpighia glabra, and Bidens pilosa, have been shown to have a number of anti-allergic properties in-vitro. The current study was conducted to determine the effects of these combined ingredients upon the nasal response to allergen challenge in patients with seasonal allergic rhinitis.     

Allelopathic properties of α-terthienyl and phenylheptatriyne,naturally occurring compounds from species of asteraceae

Alpha-terthienyl (-T), a naturally occurring polyacetylene derivative from roots ofTagetes erecta L., and phenylheptatriyne (PHT), from leaves ofBidens pilosa L., were tested as possible allelopathic agents against four seedling species (Asclepias syriaca L.,Chenopodium album L.,Phleum pratense L.,Trifolium pratense L.).Asclepias was the most sensitive of the species. Allelopathic activity was enhanced in the presence of sunlight or sources of near-UV, with LC_50s forA. syriaca of 0.15 ppm and 0.66 ppm with -T and PHT, respectively; 0.27 and 0.85 forC. album; 0.79 and 1.43 forP. pratense, and 1.93 and 1.82 forT. pratense. Near-UV exposure was saturating but never more than found in summer sunlight at Ottawa, Canada. Growth inhibition was observed with seedlings treated with -T and PHT but without near-UV irradiation. Germination of seedlings was also sensitive to -T and PHT with or without near-UV treatment. -T was extracted from soil surrounding the roots ofTagetes. Concentrations calculated for the soil (0.4 ppm) indicate that seedling growth could be significantly hindered. The activity and specificity of -T was sufficiently high to warrant future field trials to assess its potential as a natural weed-control agent.     

Perspectives on allelopathy in mexican traditional agroecosystems: A case study in tlaxcala

Agroecosystems in Tlaxcala, Mexico, are surrounded by trees and water channels and have a great variety of cultivated and noncultivated plants. The main results of a study carried out on a traditional agroecosystem in Santa Inés, Tlaxcala are presented. Some ecological aspects of polycultures, plant covers (dry leaves ofAlnus firmifolia, Berula erecta, andJuncus sp.), and the allelopathic potential of crops and noncultivated plants (fresh and dry material) were analyzed. The main plants (trees, shrubs, and herbs) present in the agroecosystem were identified. The total number of weeds in plots where plant covers were added was reduced. The number of nodules ofRhizobium phaseoli and the production of bean and squash increased with plant covers. Corn, beans, and squash showed a clear allelopathic effect, as well asChenopodium murale, Tradescantia crassifolia, Melilotus indicus, andAmaranthus hybridus, among other weeds. The contribution of allelopathy in studies of traditional agroecosystems is of great importance for the management of species in space and time. Allelopathy can be the basis of biological control of pests and weeds and of the discovery of new useful substances.     

<Emphasis Type="Italic">Bidens pilosa</Emphasis> L. Exhibits High Sensitivity to Coumarin in Comparison with Three Other Weed Species

Nine natural plant compounds were screened for phytotoxicity to Bidens pilosa L. a troublesome weed in field and plantation crops. The sensitivity of three other weed species to coumarin, the most active identified compound, was also evaluated. Coumarin, at a concentration of 500 μM, had little effect on germination and growth of Senna obtusifolia L., Euphorbia heterophylla L., and Ipomoea grandifolia L. when compared with its effects on B. pilosa L. In a concentration range of 10–100 μM, coumarin caused a dose-dependent inhibition of germination and growth of B. pilosa L. The measurements of some parameters of energy metabolism revealed that coumarin-treated root tissues exhibited characteristics of seedlings in an earlier stage of growth, including higher respiratory activity and higher activities of alcohol dehydrogenase and lipoxygenase. These results suggest that coumarin inhibition of germination and growth of B. pilosa L. was not a consequence of an impairment of energy metabolism. Rather, it seems to act as a cytostatic agent, retarding germination. At concentrations above 50 μM, coumarin increased lipoxygenase activity and the level of conjugated dienes of root extracts, suggesting that it may induce oxidative stress in seedling roots.     

Phytotoxic and antimicrobial activity of volatile constituents ofArtemisia princeps var.orientalis

The volatile constituents ofArtemisia princeps var.orientalis (wormwood) were investigated for phytotoxic and antimicrobial activities. The germination and radicle elongation of receptor plants were inhibited by volatile substances emitted from wormwood leaf and effects were concentration-dependent. Essential oil of the plant extracted by Karlsruker's apparatus suppressed seed germination and seedling elongation of the receptor plants at a threshold concentration of 4.8l/100 ml.Escherichia coli was not susceptible to the wormwood essential oil, but the growth ofBacillus subtilis, Aspergillus nidulans, Fusarium solani, andPleurotus ostreatus was inhibited severely.     

Comparative study of allelopathy as exhibited byProsopis Juliflora swartz andProsopis cineraria (L) druce

The allelopathic effects ofProsopis juliflora were studied both in the laboratory and in nature and compared with that ofProsopis cineraria to understand the chemical nature of allelochemics. Both species occupy the same habitats butP. cineraria does not appear to have any toxic effect on other plants under its canopy.P. juliflora is highly allelopathic and does not allow the growth of any other species. Leaf extracts and leaf leachates ofP. juliflora were inhibitory. Decaying leaves were also inhibitory at early stages of decomposition. Live roots were not found to be inhibitory in cogermination and interplanting of seeds. Chemical investigation of the extracts showed the allelopathic compounds to be phenolic in nature in both the species. Slow decomposition and heavy accumulation of leaf litter belowP. juliflora may possibly result in accumulation of toxic substances in soil layers, inhibiting growth of other species.     

Allelopathic Potential of Acacia confusa and Related Species in Taiwan

Acacia confusa (an endemic species) and other introduced species, namely A. aulacocarpa, A. auricumiformis, A. cincinnata, A. crassicarpa, A. leptocarpa, A. margium, A. polystachya, and A. torfilis were evaluated for allelopathic potential. Among these, A. confusa is widely distributed on the hills and lowlands of Taiwn and often exhibits a unique pattern of weed exclusion under stands. Four study sites were selected. Field observations and measurements were carried out at sites that exhibited relatively pure stands of A. confusa. Although the diversity of understory species was comparatively higher in Acacia stands than in adjacent weedy sites, the total coverage and biomass of understory plants was significantly lower than in control sites. Aqueous extracts (0.5%, 1%, 2%, 3%, 4%, and 5%) of dry leaves and litter of Acacia confusa and other species collected from various sites and dates were bioassayed using lettuce, alfalfa, and Chinese cabbage to examine their phytotoxicity. Most extracts exhibited phytotoxicity even at a concentration as low as 0.5%. Inhibition of radicle growth of test plants varied with sampling sites and dates. Bioassay with 5% extracts or above produced more than 85% inhibition of test plants regardless of habitat. In addition, surface soils collected from the upper 20 cm layer of Acacia stands revealed significant inhibition as compared with adjacent grassland control soils. The most inhibitory compounds isolated from the ether fraction of aqueous extracts were identified as: ferulic, vanillic, caffeic, gallic, m-hydroxybenzoic, and m-hydroxyphenylacetic acids. Unidentified flavonoids were also found. Chromatographic bioassays of compounds isolated from both ethyl acetate and water fractions of methanolic extracts of Acacia leaves also showed significant phytotoxicity but none was found in the fractions of chloroform and hexane, suggesting the phytotoxic compounds present in Acacia plants are water soluble.     

Effects of some compounds isolated fromCelaenodendron mexicanum standl (euphorbiaceae) on seeds and phytopathogenic fungi

The known compounds friedelin, maytensifolin B, ginkgetin, bilobetin, and amentoflavone as well as the new triterpene 3-hydroxyfriedelan-16-one were isolated fromCelaenodendron mexicanum (Euphorbiaceae), an endemic tree of the Pacific Ocean coast of Mexico. The compounds' structures were established on the basis of spectral analysis. The biological effects of aqueous leachates, a CHCl₃-MeOH extract and the isolated compounds of leaves and twigs were evaluated on the radicle growth of two plants,Amaranthus leucocarpus andEchinochloa crusgalli, and on the radial growth of three phytopathogenic fungi,Fusarium sp.,Helminthosporium sp., andAlternaria sp. The organic extracts of both leaves and twigs inhibitedAmaranthus and stimulatedEchinochloa radicle growth. On the contrary, friedelin and maytensifolin B stimulatedAmaranthus and inhibitedEchinochloa. The target fungi showed a different response to each treatment, from inhibition to stimulation.Taken in part from the M.Sc. thesis of P. Castañeda.On sabbatical leave from Departamento de Preparatoria Agrícola Universidad Autónoma de Chapingo, México.