Allelopathy

Casuarina equisetifolia litter completely suppresses germination of understory plants as shown here despite the relative openness of the canopy and ample rainfall (>120 cm/yr) at the location

Allelopathy is a biological phenomenon by which an organism produces one or more biochemicals that influence the 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. 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.[2][1]

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 Spotted Knapweed (Centaurea maculosa), Garlic Mustard (Alliaria petiolata), 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.

Contents

History

The term allelopathy, from the Greek words allele 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) 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 Homeopathy Society 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 B.C. noticed the inhibitory effects of pigweed on alfalfa. In China around the first century A.D. Yang and Tang described 267 plants that had pesticidal abilities, including those with allelopathic effects. The Swiss botanist De Candolle, in 1832 suggested that crop plant exudates were responsible for an agriculture problem called soil sickness.

Allelopathy has not been universally accepted among ecologists, and up to the early part of this century many ecologists argued that the effects of competition could not be distinguished from so called allelopathy. Competition is a negative affect that happens when two or more organisms attempt to directly use the same resource, allelopathy, on the other hand differs by indirectly affecting other organism after the input of substances into the environment. 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]

Examples of allelopathy

One of the most studied aspects of allelopathy is the role of allelopathy in agriculture. Current research is focused on the effects of weeds on crops, crops on weeds, and crops on crops. This research furthers the possibility of using allelochemicals as growth regulators and natural herbicides, to promote sustainable agriculture. 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.[7] 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.

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.[8] 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.[9] A detailed history of this interesting story can be found in Halsey 2004.[10]

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. Eucalyptus leaf litter and root exudates 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.[11]

Many crop cultivars show strong allelopathic properties, of which rice (Oryza sativa) has been most studied. 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.[12]

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.[13]

A study of Kochia scoparia in northern Montana by two high school students[14] 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.[15] A larger study later showed that Kochia seems to exhibit allelopathy on various crops in northern Montana.[16]

References

  1. ↑ 1.0 1.1 Stamp, Nancy (March 2003), "Out of the quagmire of plant defense hypotheses", The Quarterly Review of Biology 78 (1): 23–55, doi:10.1086/367580, PMID 12661508. 
  2. ↑ Fraenkel, Gottfried S. (May 1959), "The raison d'Etre of secondary plant substances", Science 129 (3361): 1466–1470, PMID 13658975. 
  3. ↑ 3.0 3.1 3.2 3.3 3.4 Willis, Rick J. (2007), "The History of Allelopathy", Springer: 3, ISBN 140204092X, http://www.google.com/books?id=C-nPBYjDAjYC&pg=PA3&, retrieved 2009-08-12 
  4. ↑ Roger, Manuel Joaquín Reigosa; Reigosa, Manuel J.; Pedrol, Nuria; González, Luís (2006), Allelopathy: a physiological process with ecological implications, Springer, pp. 1, ISBN 1402042795 
  5. ↑ Rice, Elroy Leon (1984), Allelopathy, (first edition, november 1974 by the same editor) (Second ed.), Academic Press, pp. 422 p, ISBN 978-0125870580 
  6. ↑ Roger, Manuel Joaquín Reigosa; Reigosa, Manuel J.; Pedrol, Nuria; González, Luís (2006), Allelopathy: a physiological process with ecological implications, Springer, pp. 2, ISBN 1402042795 
  7. ↑ Cornes, D. 2005. Callisto: a very successful maize herbicide inspired by allelochemistry. Proceedings of the Fourth World Congress on Allelopathy [1]
  8. ↑ Muller, C.H., Muller, W.H. and Haines, B.L. 1964. Volatile growth inhibitors produced by aromatic shrubs. Science 143: 471-473. [2]
  9. ↑ Bartholomew, B. 1970. Bare zone between California shrub and grassland communities: The role of animals. Science 170: 1210-1212. [3]
  10. ↑ Halsey, R.W. 2004. In search of allelopathy: An eco-historical view of the investigation of chemical inhibition in California coastal sage scrub and chamise chaparral. Journal of the Torrey Botanical Society 131: 343-367. The California Chaparral Institute also offers a PDF-format version of this paper. [4]
  11. ↑ Khanh, T.D, Hong, N.H., Xuan, T.D. Chung, I.M. 2005. Paddy weed control by medical and leguminous plants from Southeast Asia .Crop Protection [doi:10.1016/j.cropro.2004.09.020]
  12. ↑ Khanh, T.D, Xuan, T.D.and Chung, I.M.2007. Rice allelopathy and the possibility for weed management. Annals of Applied Biology [doi:10.1111/j.1744-7348.2007.00183.x]
  13. ↑ Stinson, K.A., Campbell, S.A., Powell, J.R., Wolfe, B.E., Callaway, R.M., Thelen, G.C., Hallett, S.G., Prati, D., and Klironomos, J.N. 2006. Invasive plant suppresses the growth of native tree seedlings by disrupting belowground mutualisms. PLoS Biology [5]
  14. ↑ For their work in this area, Overcast & Cox were awarded a first place team prize at the International Science and Engineering Fair (ISEF) in 2001.
  15. ↑ M.C. Overcast, J.J. Brimhall. 2000. Allelopathic Effects of Selected Weed Exudates on Germination and Early Growth of Triticum aestivum in Northern Toole County, Montana. [6]
  16. ↑ M.C. Overcast, D.R. Cox. 2001. Effects of Allelochemicals Produced by Kochia scoparia on Selected Crops Grown in North Toole County (NTC), Montana.

Further reading

External links