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46<br />

ANA LUISA ANAYA<br />

to assist with molecular studies conducted in today’s expanding field. Such instrumental<br />

techniques are herein recommended as economically efficient means of advancing<br />

the rigor of allelopathy research and assisting the development of a better understanding<br />

of the chemical basis for the allelopathy phenomenon (Haig, 2001).<br />

4. THE MODE OF ACTION OF ALLELOCHEMICALS<br />

Allelopathic chemicals alter plant growth and development by a multiplicity of actions<br />

on physiological processes because there are hundreds of different structures and many<br />

of the compounds have several phytotoxic effects. Whole plants bioassays and<br />

physiological tests designed to use a small quantity of compound are keys to strategies<br />

for elucidating mechanisms of action. Insight into the action of the responsible<br />

chemicals is critical to a more complete explanation of allelopathy and to its application<br />

for improving crop production. Unfortunately, we still find that linkages between<br />

inhibition of growth and the corresponding physiological mechanism are elusive. A<br />

major part of the problem is the array and diversity of allelochemicals. Several hundred<br />

different compounds have been identified, and we speculate that many others will be<br />

eventually being discovered. Most instances of allelopathic inhibition are the result of<br />

the simultaneous action of several compounds, and often these include compounds<br />

whose chemistry is divergent (Einhellig, 2002). The visible evidence in bioassays is<br />

that many phenolics, quinones, sesquiterpene lactones, alkaloids, and others alter<br />

root morphology. Although the cell membrane is an early interface with<br />

allelochemicals, relatively little attention has been given to membrane-related effects<br />

and their molecular targets.<br />

Effects of a single allelochemical or their mixtures on physiological processes<br />

include disruption of membrane permeability (Galindo, et al., 1999), ion uptake (Yu<br />

and Matsui, 1997), inhibition of electron transport in both the photosynthesis and<br />

respiratory chains (Calera et al., 1995a; Abrahim, et al., 2000), alteration of enzymatic<br />

activity (Politycka, 1999, Romagni, et al., 2000), and inhibition of cell division (Cruz-<br />

Ortega et al., 1988; Anaya and Pelayo-Benavides, 1997). In other examples, bioactivitydirected<br />

fractionation of the methanol extract of the roots of Ratibida mexicana resulted<br />

in the isolation of two bioactive sesquiterpene lactones, isoalloalantolactone and elema-<br />

1,3,11-trien-8,12-olide. Both compounds caused a significant inhibition of the radicle<br />

growth of Amaranthus hypochondriacus and Echinochloa crus-galli, exerted moderate<br />

cytotoxicity activity against three different solid tumour cell lines and inhibited the<br />

radial growth of three phytopathogenic fungi. Isoalloalantolactone also caused the<br />

inhibition of ATP synthesis, proton uptake, and electron transport (basal,<br />

phosphorylating and uncoupled) from water to methylviologen therefore acting as a<br />

Hill’s reaction inhibitor.The lactone inhibited only photosystem II (Calera et al., 1995b).<br />

Other compounds studied by Calera et al., (1995c) were the resin glycoside mixture<br />

from Ipomoea tricolor. They tested its effect on seedling growth and plasma membrane<br />

H + -ATPase activity in Echinochloa crus-galli. The resin glycoside mixture as well as<br />

Tricolorin A, the main compound in the mixture, inhibited the activity of the plasma<br />

membrane ATPase. In other studies, Cruz-Ortega et al. (1998) observed plasma

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