Weeds are known to cause enormous losses due to their interference in agroecosystems. Because of environmental and human health concerns, worldwide efforts are being made to reduce the heavy reliance on synthetic herbicides that are used to control weeds. In this regard the phenomenon of allelopathy, which is expressed through the release of chemicals by a plant, has been suggested to be one of the possible alternatives for achieving sustainable weed management. The use of allelopathy for controlling weeds could be either through directly utilizing natural allelopathic interactions, particularly of crop plants, or by using allelochemicals as natural herbicides. In the former case, a number of crop plants with allelopathic potential can be used as cover, smother, and green manure crops for managing weeds by making desired manipulations in the cultural practices and cropping patterns. These can be suitably rotated or intercropped with main crops to manage the target weeds (including parasitic ones) selectively. Even the crop mulch/residues can also give desirable benefits. Not only the terrestrial weeds, even allelopathy can be suitably manipulated for the management of aquatic weeds. The allelochemicals present in the higher plants as well as in the microbes can be directly used for weed management on the pattern of herbicides. Their bioefficacy can be enhanced by structural changes or the synthesis of chemical analogues based on them. Further, in order to enhance the potential of allelopathic crops, several improvements can be made with the use of biotechnology or genomics and proteomics. In this context either the production of allelochemicals can be enhanced or the transgenics with foreign genes encoding for a particular weed-suppressing allelochemical could be produced. In the former, both conventional breeding and molecular genetical techniques are useful. However, with conventional breeding being slow and difficult, more emphasis is laid on the use of modern techniques such as molecular markers and the selection aided by them. Although the progress in this regard is slow, nevertheless some promising results are coming and more are expected in future. This review attempts to discuss all these aspects of allelopathy for the sustainable management of weeds.

Allelopathic Interactions and Allelochemicals: New Possibilities for Sustainable Weed Management

Effects of alfalfa leaf extracts and phenolic allelochemicals on early seedling growth and root morphology of alfalfa and barnyard grass

The basis for differential allelopathic potentials among sorghum (Sorghum bicolor L. Moench) hybrids was investigated by conducting quantitative and qualitative studies of their phenolic contents. Total phenolic content in sorghum plant parts varied within hybrids, among hybrids, and between growing seasons. Inhibition of wheat (Triticum aestivum L.) radicle growth was positively associated (r=0.66) with concentrations of total phenolics contained in plant parts. Extracts from culms contributed the higherst proportion of toxicity from sorghum plants, inhibiting radicle growth up to 74.7%. Concentrations of five phenolic acids,p-hydroxybenzoic (POH), vanillic (VAN), syringic (SYR),p-coumaric (PCO), and ferulic (FER), differed in all plant parts of the three sorghum hybrids. Concentrations of POH, VAN, and SYR were consistently higher than PCO and FER. PCO and FER wer absent from some plant parts, with FER being the most frequently missing. Inhibition of wheat radicle growth was found to be positively associated with the concentration of each phenolic acid. Vanillic acid was most highly associated (r=0.44) with inhition. Thus, above-ground sorghum tissues contained phenolic acids that contributed to allelopathic potential. Additionally, sorghum roots exuded POH, VAN, and SYR that may enhance the overall allelopathic potential of sorghum during growth and after harvest when residues remain on the soil surface or are incorporated prior to planting a subsquent crop.

A chemical basis for differential allelopathic potential of sorghum hybrids on wheat

Bulk-soil and rhizosphere bacteria are thought to exert considerable influence over the types and concentrations of phytotoxins, including phenolic acids, that reach a root surface. Induction and/or selection of phenolic acid-utilizing (PAU) bacteria within the bulk-soil and rhizosphere have been observed when soils are enriched with individual phenolic acids at concentrations ≥0.25 μmol/g soil. However, since field soils frequently contain individual phenolic acids at concentrations well below 0.1 μmol/g soil, the actual importance of such induction and/or selection remains uncertain. Common bacteriological techniques (e.g., isolation on selective media, and plate dilution frequency technique) were used to demonstrate in Cecil Ap soil systems: (1) that PAU bacterial communities in the bulk soil and the rhizosphere of cucumber seedlings were induced and/or selected by mixtures composed of individual phenolic acids at concentrations well below 0.25 μmol/g soil; (2) that readily available carbon sources other than phenolic acids, such as glucose, did not modify induction and/or selection of PAU bacteria; (3) that the resulting bacterial communities readily utilize mixtures of phenolic acids as a carbon source; and (4) that depending on conditions (e.g., initial PAU bacterial populations, and phenolic acid concentration) there were significant inverse relationships between PAU bacteria in the rhizosphere of cucumber seedlings and absolute rates of leaf expansion and/or shoot biomass. The decline in seedling growth could not be attributed to resource competition (e.g., nitrogen) between the seedlings and the PAU bacteria in these studies. The induced and/or selected rhizosphere PAU bacteria, however, reduced the magnitude of growth inhibition by phenolic acid mixtures. For a 0.6 μmol/g soil equimolar phenolic acid mixture composed of p-coumaric acid, ferulic acid, p-hydroxybenzoic acid, and vanillic acid, modeling indicated that an increase of 500% in rhizosphere PAU bacteria would lead to an approximate 5% decrease (e.g., 20–25%) in inhibition of absolute rates of leaf expansion. As far as we know, this is the first time that such a relationship has been quantified.

Induction and/or selection of phenolic acid-utilizing bulk-soil and rhizosphere bacteria and their influence on phenolic acid phytotoxicity.

An investigation of the chemical basis for rice allelopathy to the rice weed arrowhead (Sagittaria montevidensis) was undertaken using GC/MS and GC/MS/MS techniques. Twenty-five compounds were isolated and identified from the root exudates of both allelopathic and non-allelopathic rice varieties. Phenolics, phenylalkanoic acids, and indoles were among the chemical classes identified. Two indoles previously unreported in rice were detected in the exudates, 5-hydroxy-2-indolecarboxylic acid and 5-hydroxyindole-3-acetic acid. Several other compounds identified in this study have not previously been reported in rice root exudates, namely mercaptoacetic acid, 4-hydroxyphenylacetic acid, and 4-vinylphenol. The levels of 15 compounds present in the exudates were quantified using GC/MS/MS. Six of the seven most abundant compounds were phenolic acids. Significant differences exist between the allelopathic and non-allelopathic cultivars in their production of three of these six compounds. Greater amounts of trans-ferulic acid, p-hydroxybenzoic acid, and caffeic acid were detected in the exudates of allelopathic cultivars. The seventh compound, abietic acid, was significantly higher in the non-allelopathic cultivars.

Identification and quantitation of compounds in a series of allelopathic and non-allelopathic rice root exudates.

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 0.015∶1 (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.

Amal A. Abdul-Rahman

Papers

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