c01 aristotletheophrastus 1991

8/7/2019 C01 AristotleTheophrastus 1991

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THEOPHRASTUS ON PI-ANT-ANIMAL INTERACTIONS

COSTAS A. TT{ANOSInstitute of General Botany, University of Athens,Athens 15784,Greece

Theophrastus,the Ancient Greekphilosopher,was born around370 BC atthe town of Eresus,in Lesbosisland and lived to the age of 85. During hislifetime the City-State system of ClassicalGreece declined and was eventuallyreplaced by a more or less unified Greek State dominated by the northernGreeks, the Macedonians,under the leadership of Alexander the Great.Theophrastus,though not a major philosopher,proved to be an extraordinaryscientist. His two voluminous works IIEPI OYTON ETOPIA: (HistoriaPlantarum, Enquiry into Plants) and IIEPI OYTON AITION (De CausisPlantarum, Causesof Plants) are the first truly scientific botanical writings andhaveled to his international recognition as the founder of the Science of Botany(e.g. Morton 1981, Evenari 1984). These works were written in their finalversion around 300 BC, exactly23 centuries ago, and their translation intoEnglish was completed only very recently.During a greatpart of his [fe, perhapsfor aslong as30 years,Theophrastuswas very closely associatedto Aristotle, whom he succeededas the director ofthe Lyceum of Athens. Aristotle, probably the maj or Ancient Greek philosopher,is further credited nowadays as the founder of Biology in general and ofZooIog in particular. It seems that Aristotle had not studied animalssystematically until 345 BC when he moved to Lesbos, together withTheophrastusand probably after the latter's suggestion.This date may constitutethe landmark of the creation of Zoolory and Botany as a result of the twofriends' joint decision to pursue an organised study of the living world.Therefore, a rough division of labour was mutually agreed and this may alsoexplain why both refrained from penetrating, in their writings, into each other'sspecific field. Nevertheless,overlapping and casual references of interactionsbetween plants and animals were obviously inevitable.Theophrastus, in particular, is repeatedly attempting comparisons andanalogies between plants and animals (the latter serving always for reference)though only in basic or general issues;for instance in discussinggeneral formand basic functions such as nutrition and reproduction. In introducing thesesubjects, Theophrastus writes: "We should not expect to find in plants acompletecorrespondencewith animals".Among severalparticular casesone can

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mention the similarity between the nature of seedsand that of eggs as well asthe analory of the leaf sheddinghabit in plantsto sheddingof horns,feathersand hair in animals.The principledcomparisonsbetweenanimalsandplantsinboth Aristotle's and Theophrastus's works reflect their conception of a singlenatural kingdom of living organisms, a profoundly scientific view derived fromthe naturalism of Ionian philosophersand in marked contrast to the idealismof Plato.

Although Theophrastus was ernphatically teaching botany instead ofproviding a local flora or a treatise on agriculture, the great economicimportance agriculture had alreadygainedduring his time is obviously reflectedto a certain degree in his works. Therefore, in numerouspassagesone findsdiscussionsabout manuring, particularly concerningthe effect of the variousanimal sourcesof manure to thegrowth of plantsof variousimportant crops.Inaddition, several referencesto animal pests and animal-causeddiseasesofplants, particularly crops, are made.Considerableattention is devoted to seedsthat are consumedby the larvae of beetles(thoughtto be producedby the seeditsel|, a mostcommoncasein legumeseedswhich are infestedby the larvae ofbeetles(today identified as bruchidbeetles).As an obviousresult of the greatimportance of galls as a tannin source, Theophrastusdealswith them in detailand describes ten different types of galls produced by oaks (Ouercus spp.), aswell as additional ones occurring in terebinth (Pistacia terebinthus) and elm(IJlmus glabra). Although the general belief of the tirne was that these wereformations of the host plant,Theophrastushadnoticed the presenceof insectswithin certain galls, as for example in the terebinth one and in the transparent,watery gall of the leaf rib of oak.

In discussingherbivory,Theophrastusmakesa strongpoint that no generalrule can be reached.Someparts of the plant may be edible and other inedible;also some animals seem to prefer the tender parts while other prefer the dryones.IIe further statesthat it is most usualthat leavesare not ediblewhile thefruits of the same plant can be consumedby both humans and animals. Lessusual is the case of lime (Tilia europaea) with edible leaves but inedible fruits.Another interesting observationis that animals find legumes a pleasure todigest. Theophrastus also mentions several examples of specific plants thatproduce toxic compounds and which may causepoisoning or death to theanimals that might consume the particular plant parts. Examples furnishedinclude black hellebore (Helleborus cl'cloph]'11us),fatal to horses and cattle, thedeadlyroot of Aconitum sp.which is not touchedby sheepor other animalsandEuonvmus spp. leaf and fruiq both fatal to sheepand especiallyto goats.Aninteresting exampleis the highlypoisonousroot of Thapsiaearganicawhich wasnever touched by the indigenous cattle of Attica, where this plant wasparticularly abundant.Importedcattle, on the other hand,might feed on it andperish of diarrhoea. The Cretan dittany (Qfiga4lq dictamnus) is an example


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of healingaction.The plant is describedby Theophrastusasbeing rare and withvery narrow distribution attributed to the fact that the goats are fond of it andgraze it down. Theophrastus adds also, somewhatsceptically though, the storyof the arrow (cited by Aristotle as well) according to which a wounded goatseeks to eat dittany and as a result the arrow drops off. This impressive storyinspired the Flemish engraver Dapper, who illustrated it in 1703 (Baumann,1982).A final note concernsthe insectrepellantaction of certain plantsor plantproducts such as olive oil, the pungent smell of marjoram (Qftganum spp.) andthe likes, as well as both speciesof Inula (i.e. eraveolensand viscosa).

In the field of pollination one hasto admit that no great contribution to ourknowledgecould be expectedduring Theophrastusera. The real nature offlowers as sexualorganseludedTheophrastusbut this is no surpriseat all sinceplant sex was suggested only two millennia later, in 1672, by the Englishphysician Thomas Millington while the first experimental proof was furnishedsubsequently,in L694, by Rudolf Jacob Camerer, a German professor ofmedicine at Tubingen(Bristow 1980).Nwertheless,Theophrastuswas able todistinguish the two sexes in certain dioecious plants on the basis of theirfruiting abfity, the female being of course the fruit bearing type.The case of thedate palm (Phqgail dactvlifera) has been well known since at least 1500 BC, asillustrated in the bas-reliefsof Nimrud in Mesopotamia.Theophrastusproceedsfurther to the point of comparing the dusting of the female flowers with themale inflorescence to what was observedwith fish, when the male sprinkles hismilt on the eggs as they are laid. Another most interesting case is thecaprification of the fig fig tree (Ficus carica) being a close companion of manfor many millennia in Eastern Mediterranean. In certain cultivated fig varietiescaprification is the necessaryprocedureto promote proper maturation of thesyconium, the complex fruit of fig. Wild figs were hung on the cultivated fig treeor wild trees were planted on eminencesnear the fig orchard to ensuretheinsectsan easyflight down wind. For it is the insectsthat after havinggrowninside the developing seeds of the wild figs will pierce the pericarp tissues,sometimesleaving a leg or a wing behind, to seekanother fig to get in andspend the rest of their life (having carried the pollen from the first to the latterwhile laying at the same time their eggs).Theseinsects are the "psenes"ofTheophrastus,the now called fig wasps,Agaontidae (the particular one beingBlastophagap!g!gc). Theophrastusalso describesanother qpe of wasps whichnever get out of the fig and are sluggishlike drones.These are actually thewingless male waspsthat fertilise the females before the latter venture their tripto their secondfig-host.

In the field of plant dispersal, numerous accounts are furnished byTheophrastus.The cormlets of the corn-flag(Gladiolussegetum)are found inmoles' runs for this animal likes them and collects them, this habit leadingunintentionallv to the dispersal of the plant. A similar case is the cachine of


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acorns by jays and other birds, resulting again in the accidentaldispersal ofoaks. An exampleof epizoochory is the fruit of Galium aoarine which sticks onthe clothes;endozoochoryis representedby ivy fruits reported to be consumedby birds. The fruits (legumes) of Colutea arborescensare described as beingeaten by sheep and their seedsare found germinating promptly from within thedroppings.A final fine exampleof zoochory is the case of the mistletoe. It isalso a masterly treatise by Theophrastuswho having identified the two speciesoccurringin Greece(Loranthuseuropaeusand Viscumalbum),asksquestionsand provides answersconcerning the peculiar habit of theseplants to grow onlyon a host tree (that may belong to a number of speciesthough).He concludesthat it is the birds that consumethe mistletoeberries;the seedspassunharmedthrough the digestive track of the bird and are able to establish their newseedlingsonly when the droppingshappento fall on a host plant.

A final point to be born in mind is that althoughTheophrastuswas by allstandards a great scholar and scientist, his works reflect precisely the overallattitude towards nature as well as the level of accumulatedknowledge of theGreek Society in general, during the end of ClassicalAntiquity.

REFERENCESBaumann H., 1982. Die Griechische Pflanzenwelt in Mlthos. Kunst undLiteratur. Hirmer Verlag, Munchen.Bristow A., 1980.The SexLife of Plants.New English Library London.Evenari M., 1984.Seedphysiolog: its history from Antiquity to the beginningof the 20th century.The BotanicalReview50:119-142.Morton A.G., 1981.Historv of Botanical Science.Academic Press,London.Theoohrastus.Enouirv into Plants.Vols 1-2.Hort A.F. ftranslator).1916and1926. William Heinemann Ltd, London and Harvard UniversiW Press.Cambridge,Mass.

Theophrastus.De Causis Plantarum.Vols 1-3.Einarson B. and Link G.K.K.(translators).7976 and 1990.William Heinemann Ltd, London and HarvardUniversity Press,Cambridge,Mass.

c01 aristotletheophrastus 1991

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