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 research of subtropical vegetation in Taiwan. IV

The phytotoxicity of plant leachates was evaluated from four subtropical grasses:Brachiaria mutica, Digitaria decumbens, Imperata cylindrica var. Major, andPanicum repens. The aqueous leachate of each grass was used to water the growth of the four grasses in pots. The leachates exhibited variable inhibition of grass growth as compared to the tap water control. By the 41st day after treatment, the leachate ofD. decumbens significantly suppressed the growth of itself and retarded that ofB. mutica andP. repens. The growth ofB. mutica was inhibited by its own leachate, but that ofI. cylindrica was not affected by any of the grass leachates. In crop growth rate (CGR) analysis, the four grass leachates exhibited a similar inhibition pattern. In laboratory bioassays, the leachates showed a significant phytotoxic effect on the radicle growth of ryegrass and lettuce. Six phytotoxic phenolics were quantitatively compared by high-performance liquid chromatography, and the amount of compounds varied with species. The highest total amount of phytotoxic phenolics occurred inD. decumbens, followed, in decreasing order, byP. repens, B. mutica, andI. cylindrica. These findings show that the leachates of four grasses possess phytotoxic compounds that may play a significant role in grass dominance in the field.     

Allelopathic influence ofSorghum bicolor on weeds during germination and early development of seedlings

The allelopathic interaction between sorghum [Sorghum bicolor (L.) Moench] and 10 species of grass and broadleaf weeds was investigated. Germination of weed seeds was slightly inhibited or stimulated, depending on species, when incubated in closed Petri dishes with germinating sorghum. Subsequent radicle and hypocotyl or coleoptile elongation of weeds was significantly inhibited by the germinating sorghum. For weeds interplanted with sorghum and grown under greenhouse conditions. The inhibitory effect on some weed species was still evident after 2 months of growth. Significant differences were found in the dry matter per weed plant grown in pots in proximity to sorghum vs. weeds grown in monoculture. Aqueous leachates from pots planted with sorghum alone or from a system in which sorghum roots protruded into water had strong allelopathic activity. These results indicate that water-soluble allelochemicals are produced by germinating sorghum seeds and that production of these substances continues during seedling growth.Agricultural Research Service, U.S. Department of Agriculture.     

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.     

Allelopathic Effects of Humus Phenolics on Growth and Respiration of Mycorrhizal Fungi

Allelopathy due to humus phenolics is a cause of natural regeneration deficiency in subalpine Norway spruce (Picea abies) forests. If inhibition of spruce germination and seedling growth due to allelochemicals is generally accepted, in contrast there is a lack of knowledge about phenolic effects on mycorrhizal fungi. Thus, this work tested effects of a humic solution and its naturally occurring phenolics on the growth and respiration of two mycorrhizal fungi: Hymenoscyphus ericae (symbiont of Vaccinium myrtillus, the main allelochemical-producing plant) and Hebeloma crustuliniforme (symbiont of P. abies, the target plant). Growth and respiration of H. crustuliniforme were inhibited by growth medium with the original humic solution (–6% and –30%), respectively, whereas the same humic solution did not affect growth but decreased respiration of H. ericae (–55%). When naturally occurring phenolics (same chemicals and concentrations in the original humic solution) were added to the growth medium, growth of H. crustuliniforme was not affected, whereas that of H. ericae significantly increased (+10%). We conclude that H. ericae is better adapted to the allelopathic constraints of this forest soil than H. crustuliniforme and that the dominance of V. myrtillus among understory species could be explained in this way.     

Allelopathic potential ofSalsola kali L. and its possible role in rapid disappearance of weedy stage during revegetation

Salsola kali L. is one of the species that invades during early successional stages on mine-spoil material, but tend to disappear rapidly from the weedy stage. What controls its rapid disappearance from the weedy stage becomes a logical question.Salsola was found to be an allelopathically active species, which also decreased the growth of selected associated species during its decaying process. However, leaf leachate did not play an important role in depressing the growth of associated species.Salsola was autotoxic, but its germinatoin was not inhibited by any of the isolated phytotoxins applied. The phytotoxins identified fromSalsola leaves were caffeic, ferulic, chlorogenic, isochlorogenic, and neochlorogenic acids, and quercetin. Allelopathy may play an important role in the rapid disappearance of the weedy stage, similar to what is established for old farm lands. However, further studies are desirable.Contribution No. R780402 from Project Reclamation, University of North Dakota supported by Grant No. 0264001 from Bureau of Mines, Department of the interior to Dr. M.K. Wali.The work was partially completed when the author was a visiting research associate professor at the University of North Dakota.     

Allelopathic potential of compounds isolated fromIpomoea tricolor cav. (Convolvulaceae)

The allelopathic potential ofI. tricolor, used in traditional agriculture as a weed controller, has been demonstrated by measuring the inhibitory activity of aqueous lixiviates and organic extracts of the plant material on seedling growth ofAmaranthus leucocarpus andEchinochloa crusgalli. Bioactivity-guided fractionation of the most active organic extract led to the isolation of the allelopathic principles, which turned out to be a mixture of glycosides, having jalapinolic acid as the aglycone portion glycosidically linked in the 11 position to an oligosaccharide composed of glucose, rhamnose, and fucose, which also combines with the carboxyl group of the aglycone to form a macrocyclic ester.This study was sponsored by Project PCECCNA-050954 of Consejo Nacional de Ciencia y Tecnología.     

Allelopathic substances and interactions ofDelonix regia (Boj) Raf

A unique pattern of weed exclusion was found under the canopy ofDelonix regia, which was planted in many places as an ornamental tree in the south of Taiwan. A quadrat method was employed to examine the botanical composition between the area underneathD. regia and its adjacent control grassland. The number of species and coverage of understory species were significantly lower in the area of theD. regia than that of the grassland, indicating the growth of understory species was suppressed byD. regia. A series of aqueous extracts of leaves, flowers, and twigs ofD. regia were bioassayed against three species to determine their phytotoxicity, and the results showed highest inhibition in the flowers. A water-culture experiment indicated the aqueous extract of flowers ofD. regia on two local understory species (Isachne nipponensis andCentella asiatica) inhibited growth of both species by more than 70%. The phytotoxicities of fallen leaves and flowers ofD. regia were not significantly affected by temperature. When the plant material was subjected to temperatures above 70°C, however, phytotoxicity was decreased, indicating that the allelopathic nature ofD. regia could easily be decomposed by fire. By means of paper, thin-layer, and high-performance liquid chromatography, and UV-visible spectrophotometry, responsible phytotoxins present in leaves, flowers, and twigs ofD. regia were identified as 4-hydroxybenzoic, chlorogenic, 3,4-dihydroxybenzoic, gallic, 3,4-dihydroxycinnamic, 3,5-dinitrobenzoic, and L-azetidine-2-carboxylic acids, and 3,4-dihydroxybenzaldehyde. The findings of bioassays and the number and amount of responsible allelopathic compounds found inD. regia are well correlated, thus permitting the conclusion that the exclusion of understory plants under the canopy ofD. regia trees was due primarily to the allelopathic effect of the fallen flower, leaves, and twigs of theD. regia. A possible mechanism of action is discussed.The study was supported by grants to C.H. Chou awarded by the Academia Sinica, Taipei, and the National Science Council (NSC-80 & 81-0211-BOO 1-25) of The Republic of China.     

Phenolic inhibitors inErica australis L. and in associated soil

Fifteen phenolic compounds were identified in methanolic extracts obtained from flowers, leaves, stems, and roots ofErica australis L. and the associated soil. It is possible that the compounds identified may be connected with allelopathic phenomena.     

Allelopathic polyacetylenes fromCentaurea repens (Russian knapweed)

The allelopathic weed Russian knapweed (Centaurea repens) was found to contain polyacetylenes VIII–XIV in the roots. Dose vs. response of the root length elongation against lettuce, alfalfa, barnyard grass, and red millet showed IX to be active. Closely related isomers were not active. Examination of the soil surrounding the Russian knapweed roots revealed the presence of IX in sufficient concentration to have an appreciable effect on the surrounding plant community.     

Allelopathic effects ofPolygonum aviculare L.

We previously reported thatPolygonum aviculare has a strong allelopathic action againstCynodon dactylon (L.) Pers. and other test species. Moreover, we found that several phenolic compounds appeared to be important allelochemics in this activity. We have now isolated other potential inhibitors fromP. aviculare residues and soil underPolygonum stands, and none of these occurred in soil underC. dactylon stands. GC-MS analysis revealed that these additional inhibitors are long-chain fatty acids with 14–22 carbons. Nine were identified inP. aviculare residues and seven in soil underP. aviculare. Sodium salts of all the identified fatty acids inhibited seedling growth ofC. dactylon and at least some test strains of the nitrogen-fixing bacteria,Azotobacter andRhizobium.     

Autointoxication mechanism ofOryza sativa

The phytotoxicity produced during decomposition of rice straw in soil was evaluated under both constant and changing temperature conditions. Bioassay tests showed that the aqueous extract from a soilstraw mixture after incubation at constant temperature was more than twice as phytotoxic as the extract from soil incubated alone. The phytotoxicity was highest at 20–25 ° C. Temperatures above 25 ° C enhanced rice straw decomposition and also degraded the phytotoxic substances more rapidly. After incubation of soil mixtures under changing temperature regimes in a phytotron, the phytotoxicy of the soil aqueous extracts increased in the following order: soil alone < soil + fertilizer < soil + straw < soil + straw + fertilizer. Growth inhibition of lettuce or rice seedlings was also at the highest at the temperature range of 25–30 ° C irrespective of the direction of temperature changes from either low to high or vice versa. Five phytotoxic phenolics,p-hydroxybenzoic, vanillic,p-coumaric, syringic, and ferulic acids, were obtained from both the aqueous extract and residue of the incubated soil samples and were quantitatively estimated by chromatography. The amount of phytotoxins found in various soil mixtures followed the same increasing order as that found by the seed bioassay test. Although no definite distribution pattern of the phenolics in the incubated soil samples can be attributed to temperature variations, the amount of the phenolics was likely higher in the samples incubated at 25 ° C than at either 15 ° C or 35 ° C. The quantity of toxins released during decomposition of rice straw in soil reached highest levels six weeks after incubation and gradually disappeared after twelve weeks.Paper No. 242 of the Scientific Journal Series of the Institute of Botany, Academia Sinica, Taipei, Taiwan, Republic of China. This study was supported by a project under the R.O.C.-U.S.A. Cooperative Science Program.     

Allelopathic potential of menthofuran monoterpenes fromCalamintha ashei

A reversed-phase HPLC analysis was used to separate and quantify five menthofuran monoterpenes inCalamintha ashei leaf soaks and washes. (+)-Evodone and desacetylcalaminthone were the major constituents of both soaks and washes. Concentrations of (+)-evodone and desacetylcalaminthone were as high as 0.66 and 0.74 mM, respectively, in leaf soaks. The highest concentration of monoterpenes in leaf washes obtained by misting was 0.021 mM. Aqueous solubilities of the menthofurans were determined to exceed concentrations required for growth inhibition. Bioassays of individualCalamintha monoterpenes demonstrated effects on germination as low as 0.05 mM for (+)-evodone. An equimolar mixture of desacetylcalaminthone and (+)-evodone reducedRudbeckia hirta germination by 17% at a combined concentration of 0.025 mM forLeptochloa dubia. Confirmation of allelopathic effects byCalamintha ashei will require long-term bioassays ofCalamintha menthofurans on the growth of native sandhill species under conditions comparable to the harsh environment of the Florida scrub.     

Autointoxication mechanism ofOryza sativa

Ammonium sulfate and potassium nitrate added as a top dressing and rice straw were incorporated into soil to understand their effect on the chemical nature of the paddy soil and on the growth and yield of rice plants during two successive crop seasons. Redox potential (Eh) determination indicated that the paddy soil appeared to be the reduced form in mixture with rice residues. The amounts of ammonium nitrogen (NH₄ ⁺—N) and nitrate nitrogen (NO₃ ^–—N) were significantly higher in the second crop than in the first crop, and the quantity of NH₄ ⁺—N was about 10 times greater than that of NO₃ ^–—N. The incorporation of rice straw decreased both the available nitrogen and the soil cations, Zn²⁺, Cu²⁺, Ca²⁺, Mn²⁺, and Na⁺. The quantity of Zn²⁺, Cu²⁺, Mg²⁺, and Na⁺ was significantly lower in the second crop that that in the first. The growth and yield of rice plants were significantly affected by cultural treatment; thus the ammonium sulfate dressing treatment resulted in higher yields than the potassium nitrate treatment. The NH₄ ⁺—N treatment had an antagonistic effect on the phytotoxic nature of rice straw decomposed in soil. The phytotoxicity (primarily phenolic type compounds) of aqueous extracts of soil varied between treatments and was significantly higher in the soil which had been mixed with rice straw. This finding correlated well with the higher amounts of phytotoxic plant phenolics produced by the decomposing rice residues in the soil.This study was supported by the National Science Council of the Republic of China.Paper No. 215 of the Scientific Journal Series, Institute of Botany, Academia Sinica.     

Allelopathic effects of giant foxtail on germination and radicle elongation of loblolly pine seed

Water extracts of giant foxtail grass inhibited germination and radicle elongation of loblolly pine seed in a laboratory test. Most of the toxic effects came from extracts of dried foxtail tops, with lesser amounts from fresh tops and roots. Soil texture affected the phytotoxicity of the extracts. Eight chemical compounds known to be phytotoxins inSetaria were identified in extracts of foxtail leaves but could not be found in soil samples under the leaves.     

Allelopathic potential ofErica vagans,calluna vulgaris,andDaboecia cantabrica

In laboratory tests the allelopathic potential ofErica vagans, Calluna vulgaris, andDaboecia cantabrica was determined. Aqueous extracts of flowers ofD. cantabrica and leaves ofC. vulgaris inhibit root and hypocotyl growth of red clover, the former causing 51% inhibition of germination. Intact aerial parts of the Ericaceae here studied drastically reduced the growth of red clover and 100% inhibition of germination was caused by flowers ofD. cantabrica. Inhibition of aqueous extracts remains after Chromatographic separation, and two well-defined inhibition zones may be observed. Hydrosoluble organic compounds (phenol-like compounds) could probably be responsible for the inhibitions detected.     

Allelopathic effects ofPolygonum aviculare L. I. Vegetational patterning

Polygonum aviculare was observed to spread rapidly into heavy stands ofCynodon dactylon (L.) Pers. resulting in death of the latter. This indicated a strong interference againstCynodon dactylon. Measurements of selected soil minerals and physical factors indicated that competition was probably not the chief cause of that interference. Soil collected under deadPolygonum was very inhibitory to all test species exceptSporobolus pyramidatus (Lam.) Hitchc., suggesting the presence of inhibitory compounds. Tops and roots ofPolygonum, root exudates, and leachate of the tops inhibited seed germination and seedling growth of most test species. Therefore, allelopathy apeared to be the dominant component of the interference, with competition probably accentuating its effects.Polygonum aviculare was inhibitory toGossypium barbadense L. andSorghum bicolor (L.) Moench, indicating that allelopathy is an important component of the interference byPolygonum against crop yields.     

Allelopathic potential in bilberry-spruce forests: Influence of phenolic compounds on spruce seedlings

Regeneration failure ofPicea abies in a subalpine bilberry-spruce forest was studied in relation to phenolic compounds, their occurrence and toxicity. Germination bioassays with natural leachates of bilberry (Vaccinium myrtillus) and spruce showed negative effects on root elongation of spruce seedlings. Growth bioassays on litter and humus demonstrated inhibitory effects of these organic layers.p-Hydroxyacetophenone, a spruce-specific metabolite, was isolated in spruce throughfall (10^–6 M), in water extracts of litter (between 1 and 8 µg/g dry wt) and organic layer (less than 1 µg/g dry wt) in addition to tannins and several common phenolic acids. Potential relationships between vegetation cover and phenolic pattern of the soil are discussed, since organic layers under bilberry heath exhibited higher amounts of phenolic acids and tannins than those under spruce.p-Hydroxyacetophenone and caffeic acid reduced, even at 5 × 10^–5 M, spruce seedling growth, especially root development, with additive effects for these two monomers. Autotoxicity involving spruce trees and allelopathy of understory species, mediated byp-hydroxy-acetophenone and other phenolic compounds, including tannins, deserves further attention in regeneration studies.     

Allelopathic effect of alfalfa (Medicago sativa) on bladygrass (Imperata cylindrica)

Greenhouse and laboratory experiments were conducted at the Agricultural and Water Resources Research Center Station, Baghdad, in 1985 and 1986 to investigate the possible allelopathic potential of alfalfa (Medicago saliva L.) and its decomposed residues on bladygrass (Imperata cylin-drica L. Beauv.), a noxious weed in Iraq, and to isolate, characterize, and quantify possible allelopathic agents in alfalfa residues and root exudates. Results indicated that decomposed alfalfa roots and their associated soil produced a 51–56% reduction in bladygrass seed germination. Root and shoot length of bladygrass seedlings were reduced by an average of 88%. Decayed and undecayed mixtures of alfalfa roots and soil at 0.0151 (w/w) inhibited bladygrass seedlings reproduced from rhizomes by 30 and 42%. It was found that root exudates of alfalfa seedlings caused significant reduction in shoot and root dry weights of bladygrass seedlings when alfalfa and bladygrass were grown together in nutrient culture. Caffeic, chlorogenic, isochloro-genic,p-coumaric,p-OH-benzoic, and ferulic acids were detected in alfalfa root exudates and residues. The highest amount (126 fig phenolic acids/g soil) of these compounds was found in alfalfa root residues after six months of decomposition in soil.     

Allelopathic potential ofIpomoea tricolor (Convolvulaceae) in a greenhouse experiment

The allelopathic potential ofIpomoea tricolor, a plant used in Mexican agriculture to control weeds, and tricolorin A, the major phytogrowth inhibitor present in the so-called resin glycosides of this plant, have been evaluated by testing leachates of the plant and the compound on the germination and radicle growth ofAmaranthus hypochondriacus, Echinochloa crusgalli, Senna uniflora, I. tricolor, andI. purpurea. The allelopathic potential ofI. tricolor was evaluated in a greenhouse experiment with dryI. tricolor mixed with sterile and nonsterile soil in pots.A. hypochondriacus was sown in pots containingI. tricolor, 2-chloro-4-(ethylamino)-6-(isopropylamino)-1,3,5 triazine (Gesaprim) or 1-glyphosphate, and the glyphosphate salt of isopropylamine (Faena), two different commercial herbicides used as a comparison toI. tricolor. Number and dry weights of different monocotyledonous and dicotyledonous weeds andA. hypochondriacus growing in the different treatments were measured.Ipomoea and Faena herbicide had a similar inhibitory effect on monocots.