Isolation and identification of allelochemicals and their activities and functions.

Allelopathy is the interaction between donor plants and receiver plants through allelochemicals. According to a great number of publications, allelopathy may be involved in several ecological aspects such as the formation of monospecific stands and sparse understory vegetation for certain plant species. Allelopathy also contributes to the naturalization of invasive plant species in introduced ranges. Autotoxicity is a particular type of allelopathy involving certain compounds. Many medicinal plants have been reported to show relatively high allelopathic activity. We selected plant species that show high allelopathic activity and isolated allelochemicals through the bioassay-guided purification process. More than 100 allelochemicals, including novel compounds have been identified in some medicinal and invasive plants, plants forming monospecific stands, plants with sparse understory vegetation, and plants showing autotoxicity. The allelopathic activity of benzoxazinones and related compounds was also determined.

Isolation and Identification of Allelopathic Substances from Forsythia suspensa Leaves, and Their Metabolism and Activity.

The fruit of Forsythia suspensa (Thunb.) Vahl has been used in traditional Chinese medicine as "Forsythiae fructus". The species is also grown in parks and gardens, and on streets and building lots, as an ornamental plant, but it requires pruning. In this study, the allelopathic activity and allelopathic substances in the leaves of pruned branches of F. suspensa were investigated to determine any potential application. The leaf extracts of F. suspensa showed growth inhibitory activity against three weed species; Echinochloa crus-galli, Lolium multiflorum, and Vulpia myuros. Two allelopathic substances in the extracts were isolated through the bioassay-guided purification process, and identified as (-)-matairesinol and (-)-arctigenin. (-)-Matairesinol and (-)-arctigenin, which showed significant growth inhibitory activity at concentrations greater than 0.3 mM in vitro. The inhibitory activity of (-)-arctigenin was greater than that of (-)-matairesinol. However, both compounds were more active than (+)-pinolesinol which is their precursor in the biosynthetic pathway. The investigation suggests that F. suspensa leaves are allelopathic, and (-)-matairesinol and (-)-arctigenin may contribute to the growth inhibitory activities. Therefore, the leaves of the pruned branches can be applied as a weed management strategy in some agricultural practices such as using the leaf extracts in a foliar spray and the leaves in a soil mixture, thereby reducing the dependency on synthetic herbicides in the crop cultivation and contributing to developing eco-friendly agriculture.

Allelopathy and Identification of Five Allelochemicals in the Leaves of the Aromatic Medicinal Tree Aegle marmelos (L.) Correa.

Aegle marmelos (L.) Correa is an economically and therapeutically valuable tree. It is cultivated as a fruit plant in southeast Asian countries. In this research, we investigated the allelopathy and possible allelochemicals in the leaves of A. marmelos. Aqueous methanol extracts of A. marmelos exhibited significant inhibitory effects against the growth of Lepidium sativum, Lactuca sativa, Medicago sativa, Echinochloa crusgalli, Lolium multiflorum, and Phleum pratense. Bioassay-directed chromatographic purification of the A. marmelos extracts resulted in identifying five active compounds: umbelliferone (1), trans-ferulic acid (2), (E)-4-hydroxycinnamic acid methyl ester (3), trans-cinnamic acid (4), and methyl (E)-3'-hydroxyl-4'-methoxycinnamate (5). The hypocotyl and root growth of L. sativum were considerably suppressed by these compounds. Methyl (E)-3'-hydroxyl-4'-methoxycinnamate also suppressed the coleoptile and root growth of E. crusgalli. The concentrations of these compounds, causing 50% growth reduction (I50) of L. sativum, were in the range of 74.19-785.4 µM. The findings suggest that these isolated compounds might function in the allelopathy of A. marmelos.

The Invasive Mechanisms of the Noxious Alien Plant Species Bidens pilosa.

Bidens pilosa L. is native to tropical America and has widely naturized from tropical to warm temperate regions in Europe, Africa, Asia, Australia, and North and South America. The species has infested a wide range of habitats such as grasslands, forests, wetlands, streamlines, coastal areas, pasture, plantations, agricultural fields, roadsides, and railway sides and has become a noxious invasive weed species. B. pilosa forms thick monospecific stands, quickly expands, and threatens the indigenous plant species and crop production. It is also involved in pathogen transmission as a vector. The species was reported to have (1) a high growth ability, producing several generations in a year; (2) a high achene production rate; (3) different biotypes of cypselae, differently germinating given the time and condition; (4) a high adaptative ability to various environmental conditions; (5) an ability to alter the microbial community, including mutualism with arbuscular mycorrhizal fungi; and (6) defense functions against natural enemies and allelopathy. The species produces several potential allelochemicals such as palmitic acid, p-coumaric acid, caffeic acid, ferulic acid, p-hydroxybenzoic acid, vanillic acid, salycilic acid, quercetin, α-pinene, and limonene and compounds involved in the defense functions such as 1-phenylhepta-1,3,5-trine, 5-phenyl-2-(1-propynyl)-thiophene, 5-actoxy-2-phenylethinyl-thiophene, and icthyothereol acetate. These characteristics of B. pilosa may contribute to the naturalization and invasiveness of the species in the introduced ranges. This is the first review article focusing on the invasive mechanisms of the species.

Phytotoxicity and Phytotoxic Substances in Calamus tenuis Roxb.

Calamus tenuis is a shrub species distributed across South Asia. It grows well in diversified habitats and tends to dominate plants in the surrounding environment. The phytotoxicity of C. tenuis and the action of its phytochemicals against other plant species could explain its dominant behavior. Compounds with phytotoxic activity are in high demand as prospective sources of ecofriendly bioherbicides. Therefore, we investigated the phytotoxicity of C. tenuis. Aqueous methanol extracts of this plant species significantly limited the growth of four test plant species, two monocots (barnyard grass and timothy), and two dicots (alfalfa and cress), in a dose- and species-dependent manner. Bio-directed chromatographic isolation of the C. tenuis extracts yielded two major active substances: a novel compound, calamulactone {(S)-methyl 8-(5-oxo-2,5-dihydrofuran-2-yl) octanoate}, and 3-oxo-α-ionone. Both of the identified compounds exerted strong growth inhibitory effects on cress and timothy seedlings. The concentrations of 3-oxo-α-ionone and calamulactone required to limit the growth of the cress seedlings by 50% (I50) were 281.6-199.5 and 141.1-105.5 µM, respectively, indicating that the effect of calamulactone was stronger with lower I50 values. Similarly, the seedlings of timothy also showed a considerably higher sensitivity to calamulactone (I50: 40.5-84.4 µM) than to 3-oxo-α-ionone (I50: 107.8-144.7 µM). The findings indicated that the leaves of C. tenuis have marked growth-inhibitory potential, and could affect surrounding plants to exert dominance over the surrounding plant community. Moreover, the two identified phytotoxic substances might play a key role in the phytotoxicity of C. tenuis, and could be a template for bioherbicide development. This paper was the first to report calamulactone and its phytotoxicity.

Allelopathic Activity of a Novel Compound, Two Known Sesquiterpenes, and a C13Nor-Isopenoid from the Leave of Croton oblongifolius Roxb. for Weed Control.

Investigation of allelopathic substances from herbal plants may lead to the development of allelochemical-based natural herbicides. Croton oblongifolius (Roxb.) is a well-known herbal plant with a long history of being used for traditional medicines and for being the source of a diverse range of bioactive compounds. This plant has been reported to have allelopathic potential; however, its allelopathic-related substances have not yet been described. Therefore, we conducted this investigation to explore the allelopathic substances from the leaves of C. oblongifolius. Aqueous methanol extracts of C. oblongifolius leaves exhibited significant growth inhibitory potential against four test plants (monocot barnyard grass and timothy, and dicot cress and lettuce). The leaf extracts were purified in various chromatographic steps and yielded four active compounds identified as (3R,6R,7E)-3-hydroxy-4-7-megastigmadien-9-one (I), 2-hydroxy alpinolide (a novel compound) (II), alpinolide (III), and epialpinolide (IV) via an analysis of the spectral data. These identified compounds significantly restricted the seedling growth of cress. The concentration necessary for 50% growth reduction of the cress seedlings varied from 0.15 to 0.24 mM for (3R,6R,7E)-3-hydroxy-4-7-megastigmadien-9-one, 0.04 to 0.11 mM for 2-hydroxy alpinolide, 0.07 to 0.12 mM for alpinolide, and 0.09 to 0.16 mM for epialpinolide. Therefore, the leaf extracts of C. oblongifolius and the characterized compounds have the potential to be used as weed-suppressive resources for natural weed control.

Invasive Mechanisms of One of the World's Worst Alien Plant Species Mimosa pigra and Its Management.

Mimosa pigra is native to Tropical America, and it has naturalized in many other countries especially in Australia, Eastern and Southern Africa and South Asia. The species is listed in the top 100 of the world's worst invasive alien species and is listed as Least Concern in the IUCN Red List of Threatened Species. M. pigra forms very large monospecific stands in a wet-dry tropical climate with conditions such as floodplains, riverbanks, grasslands, forests and agricultural fields. The stands expand quickly and threaten the native flora and fauna in the invasive ranges. Possible mechanisms of the invasion of the species have been investigated and accumulated in the literature. The characteristics of the life history such as the high reproduction and high growth rate, vigorous mutualism with rhizobia and arbuscular mycorrhizal fungi, very few natural enemies, and allelopathy, and certain secondary metabolites may contribute to the invasiveness and naturalization of M. pigra. Herbicide application, such as aerial spraying, foliar, cut-stump and soil treatments, is the primary control methods of M. pigra. The investigation of the natural enemies of M. pigra has been conducted in its native ranges since 1979, and biological control agents have been selected based on host specificity, rearing and availability. Mechanical control practices, such as hand weeding, bulldozing, chaining and fire, were also effective. However, the species often regrow from the remaining plant parts. Integration of multiple weed control practices may be more effective than any single practice. This is the first review article focusing on the invasive mechanism of M. pigra.

The Impact and Invasive Mechanisms of Pueraria montana var. lobata, One of the World's Worst Alien Species.

Pueraria montana var. lobata is native to East Asia, and was introduced to many countries due to its potential for multiple uses. This species escaped under the management conditions soon after its introduction, and became a harmful weed species. This species has been listed in the top 100 of the world's worst invasive alien species. P. montana stands expand quickly and threaten the native flora and fauna including microbiota. This species affects the concentration of carbon and nitrogen in soil and aquatic environments, and increases the amount of pollutants in the local atmosphere. Its infestation also causes serious economic losses on forestry and agriculture. Its characteristics of fast growth, thick canopy structure, enormous vegetative reproduction, and adaptative ability to the various environmental conditions may contribute to the invasiveness and naturalization of this species. The characteristics of P. montana regarding their defense functions against their natural enemies and pathogens, and allelopathy may also contribute to the invasiveness of this species. Potential allelochemicals such as xanthoxins, p-coumaric acid, caffeic acid, methyl caffeate and daidzein, and two isoflavones with anti-virus activity were identified in this species. In addition, fewer herbivore insects were found in the introduced ranges. These characteristics of P. montana may be involved in the invasive mechanisms of the species. This is the first review article focusing on the invasive mechanisms of this species.

Allelopathic Activity of the Invasive Plant Polygonum chinense Linn. and Its Allelopathic Substances.

Polygonum chinense Linn., belonging to the Polygonaceae family, is distributed mostly in northern temperate climates. This species is a high-risk invasive plant and is thought to possess allelopathic potential. This study aimed to isolate and identify the allelopathic substances from P. chinense. Aqueous methanol extracts of P. chinense significantly inhibited the growth of alfalfa and Italian ryegrass seedlings in a species- and concentration-dependent manner. Activity-guided fractionation led to the isolation of two active compounds: dehydrovomifoliol and loliolide. A cress bioassay was used to determine the biological activity of dehydrovomifoliol, and cress, alfalfa, and Italian ryegrass were used to determine loliolide. Dehydrovomifoliol significantly suppressed the seedling growth of cress at the concentration of 1 mM, and the concentrations necessary for 50% growth inhibition (I50 values) of the roots and shoots were 1.2 and 2 mM, respectively. Loliolide significantly suppressed the shoot growth of cress, alfalfa, and Italian ryegrass at the concentration of 1 mM, and the concentrations necessary for I50 values of the shoots and roots were 0.15 to 2.33 and 0.33 to 2.23 mM, respectively. The findings of our study suggest the extracts of P. chinense might have growth-inhibitory potential and that dehydrovomifoliol and loliolide might contribute as allelopathic agents.

Isolation and Identification of Plant-Growth Inhibitory Constituents from Polygonum chinense Linn and Evaluation of Their Bioherbicidal Potential.

Polygonum chinense Linn. is a medicinal and invasive plant that belongs to the family Polygonaceae. The pharmacological activities and phytochemical constituents of Polygonum chinense are well reported, but the allelopathic effects and potent allelopathic substances of P. chinense remain to be investigated. Hence, this experiment was conducted to separate and characterize potentially allelopathic substances from an extract of the Polygonum chinense plant. The Polygonum chinense plant extracts highly suppressed the growth of cress (Lepidium sativum L.), lettuce (Lactuca sativa L.), barnyard grass (Echinochloa crusgalli (L.) P. Beauv.), and timothy grass (Phleum pratense L.) seedlings in a species- and concentration-dependent way. Two active substances were separated using a series of purification procedures and determined through spectral analysis as (-)-3-hydroxy-ß-ionone and (-)-3-hydroxy-7,8-dihydro-ß-ionone. These two compounds significantly suppressed the seedling growth of Lepidium sativum (cress) at concentrations of 0.01 and 1 mM, respectively. The extract concentrations necessary for 50% growth inhibition (I50 values) of the cress hypocotyls and roots were 0.05 and 0.07 mM for (-)-3-hydroxy-ß-ionone, respectively, and 0.42 and 1.29 mM for (-)-3-hydroxy-7,8-ß-ionone, respectively. These findings suggest that these two compounds are in charge of the inhibitory effects of the Polygonum chinense extract and may serve as weed control agents.

Defensive Molecules Momilactones A and B: Function, Biosynthesis, Induction and Occurrence.

Labdane-related diterpenoids, momilactones A and B were isolated and identified in rice husks in 1973 and later found in rice leaves, straws, roots, root exudate, other several Poaceae species and the moss species Calohypnum plumiforme. The functions of momilactones in rice are well documented. Momilactones in rice plants suppressed the growth of fungal pathogens, indicating the defense function against pathogen attacks. Rice plants also inhibited the growth of adjacent competitive plants through the root secretion of momilactones into their rhizosphere due to the potent growth-inhibitory activity of momilactones, indicating a function in allelopathy. Momilactone-deficient mutants of rice lost their tolerance to pathogens and allelopathic activity, which verifies the involvement of momilactones in both functions. Momilactones also showed pharmacological functions such as anti-leukemia and anti-diabetic activities. Momilactones are synthesized from geranylgeranyl diphosphate through cyclization steps, and the biosynthetic gene cluster is located on chromosome 4 of the rice genome. Pathogen attacks, biotic elicitors such as chitosan and cantharidin, and abiotic elicitors such as UV irradiation and CuCl2 elevated momilactone production through jasmonic acid-dependent and independent signaling pathways. Rice allelopathy was also elevated by jasmonic acid, UV irradiation and nutrient deficiency due to nutrient competition with neighboring plants with the increased production and secretion of momilactones. Rice allelopathic activity and the secretion of momilactones into the rice rhizosphere were also induced by either nearby Echinochloa crus-galli plants or their root exudates. Certain compounds from Echinochloa crus-galli may stimulate the production and secretion of momilactones. This article focuses on the functions, biosynthesis and induction of momilactones and their occurrence in plant species.

Allelopathic Potential of Marsdenia tenacissima (Roxb.) Moon against Four Test Plants and the Biological Activity of Its Allelopathic Novel Compound, 8-Dehydroxy-11ß-O-Acetyl-12ß-O-Tigloyl-17ß-Marsdenin.

Plant parts and extracts that are rich in bioactive substances with allelopathic potential can be explored as a possible alternative to herbicides for natural weed control in sustainable agriculture. In the present study, we investigated the allelopathic potential of Marsdenia tenacissima leaves and its active substances. Aqueous methanol extracts of M. tenacissima showed significant inhibitory activities against the growth of lettuce (Lactuca sativa L.), alfalfa (Medicago sativa L.), timothy (Phleum pratense L.), and barnyard grass (Echinochloa crusgalli (L.) Beauv.). The extracts were purified through various chromatography steps, and one active substance was isolated and determined by spectral data to be a novel compound, assigned as steroidal glycoside 3 (8-dehydroxy-11ß-O-acetyl-12ß-O-tigloyl-17ß-marsdenin). Steroidal glycoside 3 significantly inhibited the seedling growth of cress at a concentration of 0.03 mM. The concentrations needed for 50% growth inhibition of the cress shoots and roots were 0.25 and 0.03 mM, respectively. These results suggest that steroidal glycoside 3 may be responsible for the allelopathy of M. tenacissima leaves.

The Allelopathic Effects of Trewia nudiflora Leaf Extracts and Its Identified Substances.

Trewia nudiflora Linn. is a woody plant of the Euphorbiaceae family. It is well known for its use as a folk remedy, but its potential for phytotoxicity has not been explored. Therefore, this study investigated the allelopathic potential and the allelopathic substances in T. nudiflora leaves. The aqueous methanol extract of T. nudiflora was found to have a toxic effect on the plants used in the experiment. The shoot and root development of lettuce (Lactuca sativa L.) and foxtail fescue (Vulpia myuros L.) were significantly (p ≤ 0.05) reduced by the T. nudiflora extracts. The growth inhibition by the T. nudiflora extracts was proportional to the extract concentration and varied with the test plant species. The chromatographic separation of the extracts resulted in the isolation of two substances, identified as loliolide and 6,7,8-trimethoxycoumarin based on their respective spectral analyses. Both substances significantly inhibited lettuce growth at a concentration of 0.01 mM. To inhibit 50% of the growth of the lettuce, the required concentration of loliolide was 0.043 to 0.128 mM, while that of 6,7,8-trimethoxycoumarin was 0.028 to 0.032 mM. Comparing these values, the lettuce growth was more sensitive to 6,7,8-trimethoxycoumarin than loliolide, suggesting that 6,7,8-trimethoxycoumarin was more effective than loliolide. Therefore, the inhibition of the growth of the lettuce and foxtail fescue suggests that loliolide and 6,7,8-trimethoxycoumarin are responsible for the phytotoxicity of the T. nudiflora leaf extracts. Thus, the growth-inhibitory effectiveness of the T. nudiflora extracts and the identified loliolide and 6,7,8-trimethoxycoumarin may be used to develop bioherbicides that restrict the growth of weeds.

Evolution of the Secondary Metabolites in Invasive Plant Species Chromolaena odorata for the Defense and Allelopathic Functions.

Chromolaena odorata (L.) R.M. King & H. Robinson is native to tropical America, and has naturalized in many other countries in tropical Asia, Austria, and West Africa. The species often forms dense thickets and reduces the native species diversity and population in the invasive ranges. The species is also considered as a noxious weed in agriculture fields, and listed in the 100 of the world's worst invasive alien species. The characteristics of its life-history such as the seed production rate, growth pattern, and adaptative ability to the environmental conditions may contribute to the invasiveness of the species. Possible evidence of the defense capacity against the natural enemy, and the allelopathic potential against the competitive plant species for C. odorata has been accumulated in the literature over three decades. The extracts, residues, and/or rhizosphere soil of C. odorata increased the mortality of various insects and parasitic nematodes, and decreased their population. The extracts, residues, and/or rhizosphere soil of C. odorata also inhibited the germination and growth of several plant species including the indigenous plant species in the invasive ranges of C. odorata. Toxic substances, pyrrolizidine alkaloids were found in the leaves and flowers of C. odorata. These pyrrolizidine alkaloids may work as the defense agents against the natural enemies. Several potential allelochemicals such as flavonoids, phenolic acids, and terpenoids were also found in the plant extracts of C. odorata. Some of these compounds may work as allelopathic agents of C. odorata and inhibit the germination and growth of the competitive plant species. These characteristics of C. odorata for the defense function against their natural enemies such as insects and parasitic nematodes, and allelopathic potential against the competitive native plant species may contribute to the invasiveness and naturalization of C. odorata in the new habitats as invasive plant species. However, it is necessary to determine the concentration of these allelochemicals in the neighboring environment of C. odorata such as the rhizosphere soil since allelochemicals are able to work only when they are released into the neighboring environment. It is the first review article focusing on the defense function and allelopathy of C. odorata.

Allelopathic Substances of Osmanthus spp. for Developing Sustainable Agriculture.

Osmanthus fragrans Lour. has been cultivated for more than 2500 years because of the fragrance and color of the flowers. The flowers and roots have been used in tea, liquors, foods, and traditional Chinese medicine. The species contains more than 180 compounds including terpenoids, phenylpropanoids, polyphenols, flavonoids, and sterols. However, there has been limited information available on the allelopathic properties and allelopathic substances of O. fragrans. We investigated the allelopathy and allelopathic substances of O. fragrans and Osmanthus heterophyllus (G.Don) P.S. Green, as well as Osmanthus × fortunei Carrière, which is the hybrid species between O. fragrans and O. heterophyllus. The leaf extracts of O. fragrans, O. heterophyllus, and O. × fortunei suppressed the growth of cress (Lepidium sativum L.), alfalfa (Medicago sativa L.), Lolium multiflorum Lam., and Vulpia myuros (L.) C.C.Gmel with the extract concentration dependently. The extract of the hybrid species O. × fortune was the most active among the extracts. The main allelopathic substances of O. × fortunei and O. fragrans were isolated and identified as (+)-pinoresinol and 10-acetoxyligustroside, respectively. (+)-Pinoresinol was also found in the fallen leaves of O. × fortunei. Both compounds showed an allelopathic activity on the growth of cress and L. multiflorum. On the other hand, several allelopathic substances including (+)-pinoresinol may be involved in the allelopathy of O. heterophyllus. O. fragrans, O. heterophyllus, and O. × fortunei are evergreen trees. but their senescent leaves fall and cover the soil under the trees. It is possible that those allelopathic substances are liberated through the decomposition process of the leaves into their rhizosphere soil, and that they accumulate in the soil and provide a competitive advantage to the species through the inhibition of the growth of the neighboring competing plants. Therefore, the leaves of these Osmanthus species are allelopathic and potentially useful for weed management options in some agriculture settings to reduce commercial herbicide dependency for the developing sustainable agriculture systems.

Identification of four allelopathic compounds including a novel compound from Elaeocarpus floribundus Blume and determination of their allelopathic activity.

Allelopathic compounds can play a vital role in protecting the environment from pollution by synthetic herbicides. Compounds isolated from plant species with allelopathic potential can be used as natural herbicides to control weeds and help reduce environmental pollution. Elaeocarpus floribundus has been reported to contain allelopathic compounds. Aqueous methanolic extracts of the leaves of this plant showed strong growth inhibitory potential against two test species (monocotyledonous Italian ryegrass and dicotyledonous alfalfa) in plants- and dose-dependent technique. Several extensive chromatographic separations of the E. floribundus leaf extracts yielded four active compounds 1, 2, 3, and 4 (novel compound). All the identified compounds showed strong growth inhibitory potential against cress. The concentrations caused for 50% growth limitation (I50 values) of the cress seedlings were in the range 500.4-1913.1 µM. The findings indicate that the identified compounds might play a pivotal function in the allelopathic potential of E. floribundus tree. This report is the first on elaeocarpunone and its allelopathic potential.

Allelopathy and Allelochemicals of Solidago canadensis L. and S. altissima L. for Their Naturalization.

Solidago canadensis L. and Solidago altissima L. are native to North America and have naturalized many other continents including Europa and Asia. Their species is an aggressive colonizer and forms thick monospecific stands. The evidence of the allelopathy for S. canadensis and S. altissima has accumulated in the literature since the late 20th century. The root exudates, extracts, essential oil and rhizosphere soil of S. canadensis suppressed the germination, growth and the arbuscular mycorrhizal colonization of several plants, including native plant species. Allelochemicals such as fatty acids, terpenes, flavonoids, polyphenols and their related compounds were identified in the extracts and essential oil of S. canadensis. The concentrations of total phenolics, total flavonoids and total saponins in the rhizosphere soil of S. canadensis obtained from the invasive ranges were greater than those from the native ranges. Allelochemicals such as terpenes, flavonoids, polyacetylene and phenols were also identified in the extracts, essential oil and the rhizosphere soil in S. altissima. Among the identified allelochemicals of S. altissima, the cis-dehydromatricaria ester may be involved in the allelopathy considering its growth inhibitory activity and its concentration in the rhizosphere soil. Therefore, the allelopathy of S. canadensis and S. altissima may support their invasiveness, naturalization and formation of thick monospecific stands. This is the first review article focusing on the allelopathy of both of S. canadensis and S. altissima.

Allelopathy and Allelochemicals of Imperata cylindrica as an Invasive Plant Species.

Imperata cylindrica is native to Southeast Asia and East Africa and has become naturalized in humid tropics, subtropics and warmer temperate zones of the world. The species is one of the top ten worst weeds in the worlds and is listed among the world's top 100 worst invasive alien species. It is an aggressive colonizer and forms large monospecific stands in several countries. Possible evidence of the allelopathy of I. cylindrica has been accumulated in the literature over three decades. The extracts, leachates, root exudates, decomposing residues and rhizosphere soil of I. cylindrica were found to suppress the germination and growth of several plant species, including woody plant species, and to reduce their rhizobium nodulation and mycorrhizal colonization. Several allelochemicals, such as fatty acids, terpenoids, simple phenolics, benzoic acids, phenolic acids, phenolic aldehydes, phenylpropanoids, flavonoids, quinones and alkaloids, were also found in the extracts, leachates, root exudates and/or growth medium of I. cylindrica. These observations suggest that allelochemicals may be synthesized in I. cylindrica and released into the rhizosphere soil and surrounding environments either by the leachates, root exudation or decomposition process of plant parts, and certain allelochemicals may contribute to the alteration of the microbial community, including rhizobia and mycorrhizal fungi, suppressing the regeneration process of native plant species through the inhibition of their germination and growth. Therefore, the allelopathy of I. cylindrica may support its invasiveness, naturalization and formation of large monospecific stands. This is the first review article focusing on the allelopathy of I. cylindrica.

Allelopathy and Allelochemicals of Leucaenaleucocephala as an Invasive Plant Species.

Leucaena leucocephala (Lam.) de Wit is native to southern Mexico and Central America and is now naturalized in more than 130 countries. The spread of L. leucocephala is probably due to its multipurpose use such as fodder, timber, paper pulp, shade trees, and soil amendment. However, the species is listed in the world's 100 worst invasive alien species, and an aggressive colonizer. It forms dense monospecific stands and threatens native plant communities, especially in oceanic islands. Phytotoxic chemical interactions such as allelopathy have been reported to play an important role in the invasion of several invasive plant species. Possible evidence for allelopathy of L. leucocephala has also been accumulated in the literature over 30 years. The extracts, leachates, root exudates, litter, decomposing residues, and rhizosphere soil of L. leucocephala increased the mortality and suppressed the germination and growth of several plant species, including weeds and woody plants. Those observations suggest that L. leucocephala is allelopathic and contains certain allelochemicals. Those allelochemicals may release into the rhizosphere soil during decomposition process of the plant residues and root exudation. Several putative allelochemicals such as phenolic acids, flavonoids, and mimosine were identified in L. leucocephala. The species produces a large amount of mimosine and accumulates it in almost all parts of the plants, including leaves, stems, seeds, flowers, roots, and root nodules. The concentrations of mimosine in these parts were 0.11 to 6.4% of their dry weight. Mimosine showed growth inhibitory activity against several plant species, including some woody plants and invasive plants. Mimosine blocked cell division of protoplasts from Petunia hybrida hort. ex E. Vilm. between G1 and S phases, and disturbed the enzyme activity such as peroxidase, catalase, and IAA oxidase. Some of those identified compounds in L. leucocephala may be involved in its allelopathy. Therefore, the allelopathic property of L. leucocephala may support its invasive potential and formation of dense monospecific stands. However, the concentrations of mimosine, phenolic acids, and flavonoids in the vicinity of L. leucocephala, including its rhizosphere soil, have not yet been reported.

Allelopathic Properties of Lamiaceae Species: Prospects and Challenges to Use in Agriculture.

Herbicide resistance due to the increasing reliance on herbicides is a near-term challenge for the world's agriculture. This has led to a desire to develop new herbicides with a novel mode of action, to address resistance in weed species. Lamiaceae, a large dicotyledonous plant family, is very well known for the multitudinous pharmacological and toxicological properties of its member species. Moreover, many species of this family are significant for their allelopathic activity in natural and laboratory settings. Thus, plants in Lamiaceae have the potential to be sources of alternative herbicides. However, gaps in our knowledge need to be addressed prior to adopting these allelopathic activities in agriculture. Therefore, we review the existing state of knowledge about the Lamiaceae family, the reported allelopathic properties of plant extracts, and their isolated allelochemicals under laboratory, greenhouse, and field conditions. In addition, we offer a perspective on existing challenges and future opportunities for adopting the allelopathic properties of Lamiaceae plant species for green agriculture.