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Allelopathy is a biological phenomenon by which an organism produces one or more biochemicals that influence the germination, growth, survival, and reproduction of other organisms. These biochemicals are known as allelochemicals and can have beneficial (positive allelopathy) or detrimental (negative allelopathy) effects on the target organisms and the community. Allelochemicals are a subset of secondary metabolites,[1] which are not required for metabolism (i.e. growth, development and reproduction) of the allelopathic organism. Allelochemicals with negative allelopathic effects are an important part of plant defense against herbivory.[1][2]

The production of allelochemicals are affected by biotic factors such as nutrients available, and abiotic factors such as temperature and pH.

Allelopathy is characteristic of certain plants, algae, bacteria, coral, and fungi. Allelopathic interactions are an important factor in determining species distribution and abundance within plant communities, and are also thought to be important in the success of many invasive plants. For specific examples, see black crowberry (Empetrum hermaphroditum), spotted knapweed (Centaurea maculosa), garlic mustard (Alliaria petiolata), Casuarina/Allocasuarina spp., and nutsedge.

The process by which a plant acquires more of the available resources (such as nutrients, water or light) from the environment without any chemical action on the surrounding plants is called resource competition. This process is not negative allelopathy, although both processes can act together to enhance the survival rate of the plant species.

History


The term allelopathy from the Greek-derived compounds allelo- and -pathy (meaning "mutual harm" or "suffering"), was first used in 1937 by the Austrian professor Hans Molisch in the book Der Einfluss einer Pflanze auf die andere - Allelopathie (The Effect of Plants on Each Other - Allelopathy) published in German.[3] He used the term to describe biochemical interactions that inhibit the growth of neighbouring plants, by another plant.[4] In 1971, Whittaker and Feeny published a study in the journal Science, which defined allelochemicals as all chemical interactions among organisms.[3] In 1984, Elroy Leon Rice in his monograph on allelopathy enlarged the definition to include all direct positive or negative effects of a plant on another plant or on micro-organisms by the liberation of biochemicals into the natural environment.[5] Over the next ten years, the term was used by other researchers to describe broader chemical interactions between organisms, and by 1996 the International Allelopathy Society (IAS) defined allelopathy as "Any process involving secondary metabolites produced by plants, algae, bacteria and fungi that influences the growth and development of agriculture and biological systems."[6] In more recent times, plant researchers have begun to switch back to the original definition of substances that are produced by one plant that inhibit another plant.[3] Confusing the issue more, zoologists have borrowed the term to describe chemical interactions between invertebrates like corals and sponges.[3]

Long before the term allelopathy was used, people observed the negative effects that one plant could have on another. Theophrastus, who lived around 300 BC noticed the inhibitory effects of pigweed on alfalfa. In China around the first century AD, the author of Shennong Ben Cao Jing described 267 plants that had pesticidal abilities, including those with allelopathic effects.[7] In 1832, the Swiss botanist De Candolle suggested that crop plant exudates were responsible for an agriculture problem called soil sickness.

Allelopathy is not universally accepted among ecologists and many have argued that its effects cannot be distinguished from the competition which results when two (or more) organisms attempt to use the same limited resource, to the detriment of one or both. Allelopathy is a direct negative effect on one organism resulting from the input of substances into the environment by another. In the 1970s, great effort went into distinguishing competitive and allelopathic effects by some researchers, while in the 1990s others argued that the effects were often interdependent and could not readily be distinguished.[3]

However, by 1994 D. L. Liu and J. V. Lowett at the Department of Agronomy and Soil Science, University of New England in Armidale, NSW, Australia wrote two papers[8][9] in the Journal of Chemical Ecology that developed methods to separate the allelochemical effects from other competitive effects, using barley plants and inventing a process to examine the allelochemicals directly. In 1994, M-C Nilsson at the Swedish University of Agricultural Sciences in UmeƄ, showed in a field study that allelopathy exerted by Empetrum hermaphroditum reduced growth of Scots pine seedlings by c. 40%, and that below-ground resource competition by E. hermaphroditum accounted for the remaining growth reduction.[10] For this work she inserted PVC-tubes into the ground to reduce below-ground competition or added charcoal to soil surface to reduce the impact of allelopathy, as well as a treatment combining the two methods.

Application


The possible application of allelopathy in agriculture is the subject of much research.[11] Current research is focused on the effects of weeds on crops, crops on weeds, and crops on crops.[12] This research furthers the possibility of using allelochemicals as growth regulators and natural herbicides, to promote sustainable agriculture.[13] A number of such allelochemicals are commercially available or in the process of large-scale manufacture. For example, Leptospermone is a purported thermochemical in lemon bottlebrush (Callistemon citrinus). Although it was found to be too weak as a commercial herbicide, a chemical analog of it, mesotrione (tradename Callisto), was found to be effective.[14] It is sold to control broadleaf weeds in corn but also seems to be an effective control for crabgrass in lawns. Sheeja (1993) reported the allelopathic interaction of the weeds Chromolaena odorata (Eupatorium odoratum) and Lantana camara on selected major crops.

Many crop cultivars show strong allelopathic properties, of which rice (Oryza sativa) has been most studied.[15][16][17] Rice allelopathy depends on variety and origin: Japonica rice is more allelopathic than Indica and Japonica-Indica hybrid. More recently, critical review on rice allelopathy and the possibility for weed management reported that allelopathic characteristics in rice are quantitatively inherited and several allelopathy-involved traits have been identified.[18]

Many invasive plant species interfere with native plants through allelopathy.[19][20] A famous case of purported allelopathy is in desert shrubs. One of the most widely known early examples was Salvia leucophylla, because it was on the cover of the journal Science in 1964.[21] Bare zones around the shrubs were hypothesized to be caused by volatile terpenes emitted by the shrubs. However, like many allelopathy studies, it was based on artificial lab experiments and unwarranted extrapolations to natural ecosystems. In 1970, Science published a study where caging the shrubs to exclude rodents and birds allowed grass to grow in the bare zones.[22] A detailed history of this story can be found in Halsey 2004.[23]

Allelopathy has been shown to play a crucial role in forests, influencing the composition of the vegetation growth, and also provides an explanation for the patterns of forest regeneration. The black walnut (Juglans nigra) produces the allelochemical juglone, which affects some species greatly while others not at all. The leaf litter and root exudates of some Eucalyptus [24] species are allelopathic for certain soil microbes and plant species. The tree of heaven, Ailanthus altissima, produces allelochemicals in its roots that inhibit the growth of many plants. The pace of evaluating allelochemicals released by higher plants in nature has greatly accelerated, with promising results in field screening.[25]

Garlic mustard is an invasive plant species in North American temperate forests. Its success may be partly due to its excretion of an unidentified allelochemical that interferes with mutualisms between native tree roots and their mycorrhizal fungi.[26]

A study of Kochia scoparia in northern Montana by two high school students[27] showed that when Kochia precedes spring wheat (Triticum aestivum), it reduces the spring wheat's growth. Effects included delayed emergence, decreased rate of growth, decreased final height and decreased average vegetative dry weight of spring wheat plants.[28] A larger study later showed that Kochia seems to exhibit allelopathy on various crops in northern Montana.[29]

See also


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