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Chemical Influences of Other Plants (Allelopathy)*. by Michael Evenari

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Chemical Influences of Other Plants (Allelopathy)*. by Michael Evenari Chemical influences of other plants (allelopathy)*. By Michael Evenari. With 11 figures. I. lntroduction. The plant in innumerable chemical ways controls its own development from germination to flowering and the ripening of fruits and seeds. These chemical controls are "self" controls as certain substances, like growth hormones, ger­ mination inhibitors etc., produced by the plant body itself, regulate the physio­ logical functions of this self-same body. Goncentrating most of the research efforts on this problern the fact has often been overlooked that the plant is at the same time a chemical powerhouse, producing and secreting chemical substances with which it actively changes its abiotic and biotic environment. Through these secretions the plant is able to influence other plants growing in its surrounding and exerts to some extent a chemical control of its environ­ ment. In some cases, as in those of many parasites, the chemical relationship between plant and plant forms a conditio sine qua non for the fulfillment of the normal life cycle of the plant concerned. The detection of the antibiotics has deepened the interest of the botanist in the problern of the chemical influence of plant upon plant. But its great importance for the understanding of basic facts in plant sociology and plant ecology is still not yet fully recognized, although our knowledge about the subject is quite old. Already KuESTER (1909) published a paper on the "Chemische Beeinflussung der Organismen durcheinander". In 1937 MoLISCH wrote his fascinating booklet on the same subject and in accorfdance with his terminology we will define allelopathy as the influence of one plant upon another occurring under natural conditions and exerted by chemical means other than nutritional ones. Lately GRÜMMER (1955) published another book on allelopathy where the main results of modern research are summed up. In basing our discussion on this definition of allelopathy we immediately encounter some difficulties. Our definition of allelopathy excludes nutritional relations between different organisms. But in certain cases of stimulation f.i. it is very difficult to discern between a nutritional effect and one produced by stimulating substances. Allelopathy exerted through secretion of hormones and enzymes should be included in our chapter. We will only touch upon this type of allelopathy as other chapters of this encyclopaedia will deal with the same question. * B. RADEMACHER's contribution "Gegenseitige Beeinflussung höherer Pflanzen" in Vol. XI, p. 655 of this encyclopaedia dealsalso with allelopathy and compliments our chapter in many points. He for instance deals with the nutritional relations between different organ­ isms which we exclude per definitionem from our own contribution. 44* G. Melchers (ed.), Aussenfaktoren in Wachstum und Entwicklung © Springer-Verlag Berlin Heidelberg 1961 692 MICHAEL EvENARI: Chemical inßuences oi other plants (allelopathy). As we decided that one of the criteria of allelopathy is its occurrence under natural conditions a laboratory experiment alone cannot prove allelopathy. Besides the difficulty to determine what are "natural conditions" we here en­ counter another problem. Very often potentially allelopathic substances such as antibiotics are present in a certain organism. A laboratory experiment proves their antibiotic effect on certain microorganisms. But there is no proof that under natural conditions the antibiotic is of any biological importance. But as such a proof is very difficult to give the substance in question may still have an allelopathic function. Wemade it a rule to exclude cases of potential allelo­ pathy when at least no indication of allelopathy under natural conditions exists. Another difficulty lies in the complexity of "natural conditions". · When dealing j.i. with the rhizosphere effect or the mycorrhiza it is often impossible to unravel the complex conditions without the most elaborate experiments (seldom done) and to come to a clear conclusion. We naturally included here allelopathy between host and parasite and between microorganisms. In these chapters the material is not exhaustively dealt with and mostly only examples are given as otherwise we would have transgressed to much the scope of this encyclopedia. We dealt extensively with ethylene as an allelopathic agent because this is the only case of allelopathy which was thoroughly investigated. The chapter by CARR on the "Chemical influence of the environment" which follows (pp. 737-794) constitutes in many ways a complement to this chapter on allelopathy. n. Allelopathy between higher plants. A. Volatile secretions. a) Ethylene. 1. Occnrrence and identification. ELMER (1932) enclosed apples and potato tubers in the same container and found that under the influence of the apples the sprouting of the tubers was delayed and the sprouts formed were abnormal. HuELIN, (1933) SMITH and GANE (1933) and Kmn and WEST (1934) obtained the same effect in using ethylene and suggested that the gas emanating from the apples in ELMER's experimentwas ethylene. GANE (1934) isolated ethylene from apples and was the first to prove by chemical identification that plants secrete ethylene and that it is physiologically active when it comes in contact with other plants. Later ethylene was isolated from ripening bananas by NIEDERL, BRENNER and KELLEY (1938), from avocados by PRATT, YoUNG and BIALE (1948), from ripening tomato fruits by SPENCER (1956, 1958) and from leaves of Silybum marianum by PRATT (1954). Table 1 gives the amounts of ethylene liberated by different plants and plant organs. GANE (1934) estimates that one apple during its whole life-time produces about 1 ml of ethylene. In addition to the fruits mentioned BIALE, YoUNG and ÜLMSTEAD (1954) have proved by chemica] identification the emanation of ethylene by the follow­ ing fruits: Annona cherimola, Feijoa sellowiana, and Oasimiroa edulis (Sapote). Mango, Lemon and Oranges do not produce ethylene. The production of ethylene by citrus fruits reported by other authors (see f.i. MrLLER 1946) was apparently based on the fact that the fungus Penicillium digitatum, which infects citrus fruits, produces ethylene (BIALE 1940, MILLER, WINSTON and FISHER 1940, BIALE and SHEPHERD 1941, YoUNG, PRATT and Physiological effects. 693 Table 1. Oomparative ethylene production by Beveral plant organtJ. (Mter BIALE 1950 and others.) Plants Ethylene producedl Author ml/kg/24 hrs. Apple var. Worcester} 0.12 GANE (1934) Apple var. Pearmain · Apple var. Gravenstein 0.10-0.28 IIANsEN and CHRisTIANSEN (1939) Apple var. Canada gris ..... 2 Apple var. Reine des Reinettes . 11 } PHA.N-CHON·TON (1956) Avocado var. Fuerte. 0.23 PRATT, YOUNG and BIALE (1948) Banana ..... 0.0005 NIEDERL, BRENNER and KELLEY (1938) Pear var. Anjou .. 0.072-0.72 and CHRISTENSEN (1939) Pear var. BartJett. 1.32 -5.33 } HANSEN Pear var. Williams 0.7-1.8 Pru.N.CHoN-TON (1956) Silybum leaves . 0.46-0.8 PRATT (1954) Cotton plants . 1.96 HALL et al. (1957) Cotton leaves .... 0.13 HALL etal. (1957) Flowers (Iris, Dahlia) 1-4 Pru.N-CHON-TON (1956) Tornato ..... 0.01-0.19 SPENCER (1956) BIALE 1951, PHAN-C:HON-TON 1957a). Some other fungi, pathogenic to humans like Blastomyces dermatitidis also produce ethylene (NrcKERSON 1948). The pathogen Fusarium the cause of fusarium wilt in tomatoes produces ethylene in culture and in the diseased host (DIMOND and WAGGONER 1953a). !IALL's (1952a) report that oranges produce ethylene is apparently based on a faulty method. HALL et al. (1957) have shown with an improved method that cotton plants and cotton leaves produce ethylene in considerable quantities. Flowers also emanate ethylene (PHAN-CHON-TON 1956). These are all the cases in which the production of ethylene by plant organs was proven cheinically. In many other cases the emanation of volatile sub­ stances, identical with near certainty with ethylene, was tested by various biological tests, like leaf epinasty and the triple response of etiolated pea seed­ lings etc. Leaves, petals, anthers, pistils, mature and immature fruits, tubers, twigs, roots etc. were shown to produce ethylene. (For details see DENNY 1935, DENNY and MILLER 1935, MoLISCH 1937 and GRÜMMER 1955.) It is even possible that soil microorganisms produce ethylene which has an allelopathic influence on higher plants growing in that soil (HALL et al. 1957). 2. Physiologieal eHeets. oc) Maturation of fruits. Already at the beginning of the century California Citrus growers used to "eure" the fruits forcably by placing them in a tent together with burning Kerosene stoves. This treatment stimulates the colouring of the fruit. SIEVERS and TRUE (1912) proved that not the heat but the gaseous combustion products of the Kerosene were responsible for the curing effect. ÜHACE and DENNY (1924) and DENNY (1924a) suggested that ethylene was the active agent. The "United Fruit Company" in shipping bananas to the United States noticed over a period of years prior to 1917 that the ripening of the bananas occurred in "pockets" which acted as a focus of ripening for the surrounding fruit. RIDLEY (1923) found that ripe bananas produce a gas which caused the accelerating of the ripening of green bananas. This gas was ethylene which proved effective in stimulating the ripening of many plant products, as shown early by DENNY (1924a, lemons), HARVEY (1925, celery), ÜHACE and 0HURCH (1927), REGEIMBAL and HARVEY (1927, pineapples), REGEIMBAL, V ACHA and Handbuch d. Pflanzenphysiologie, Bd. XVI. 44a 694 MICHAEL EvENARI: Chemical influences
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