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Juglans Spp., Juglone and Allelopathy

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Juglans Spp., Juglone and Allelopathy AllelopathyJournatT(l) l-55 (2000) O Inrernationa,^,,r,':'r::;:';::::,:rt;SS Juglansspp., juglone and allelopathy R.J.WILLIS Schoolof Botany.L.iniversity of Melbourre,Parkville, Victoria 3052, ALrstr.alia (Receivedin revisedform : February 26.1999) CONTENTS 1. Introduction 2. HistoricalBackground 3. The Effectsof walnutson otherplants 3.i. Juglansnigra 3.1.1.Effects on cropplants 3. I .2. Eft'ectson co-plantedtrees 3. 1 .3 . Effectson naturalvegetation 3.2. Juglansregia 3.2.1. Effectson otherplalrts 3.2.2.Effects on phytoplankton 1.3. Othel walnuts : Juglans'cinerea, J. ntttlor.J. mandshw-icu 4. Juglone 5. Variability in the effect of walnut 5.1. Intraspecificand Interspecific variation 5.2. Seasonalvariation 5.3 Variation in the effect of Juglansnigra on other.plants 5.4. Soil effects 6. Discussion Ke1'rvords: Allelopathy,crops, history, Juglan.s spp., juglone. phytoplankton,walnut, soil, TTCCS 1. INTRODUCTION The"rvalnuts" are referable to Juglans,a genusof 20-25species with a naturaldistribution acrossthe Northern Hemisphere and extending into SouthAmerica. Juglans is a memberof thefamily Juglandaceae which contains6 or 7 additionalgenera including Cruv,a, Cryptocctrva and a total of about 60 species. Walnuts are corrunerciallyimportant as the sourceof the ediblewalnut, the highly prizedtimber and as a specimentrees. Eating walnutsare usually obtarnedfrom -/. regia (the colrunonor Persianwalnut, erroneousll'known as the English walnut)- a nativeof SEEurope and Asia, which haslong been cultivated, but arealso sometin.res availablelocally from other speciessuch as J. nigra (back walnut) - a native of eastern North America andJ. ntajor, J. calfornica andJ. hindsii, native to the u,esternu.S. ILillis Grafting of supcrior fnrit-bearing scions of J. regia onlo rootstocksof hlrdier spccics. strclras J' ltind'sii or.I. nigra is cornruonpractice. The speciesof ualnnts are in necd of relis-ion.particularly thosefrom South and Central America, and frorn East Asia. as more than 60 specieshave been recognisedin the past. At preseut about 20-25 speclcsand ntttnerous varieties are recognised:the better knorln speciesare listecl in Table L ihbleL Majorspccies ol Jttglart.s(contpiled lionr varioussourccs) Species Common name Distribution Europe/Asia .1.regio L. pcrsian Conrr.nor.ror rvalnut S.E .Eulope.Asia: rvidelv planted .J. ntotrd.slturica Maxinr. Manchurian rvalnut N.I:. Asia .1.oilatrrltifolirrCarriere .Iapaneses,alnut .lapan.Russia ,J. cotha|ert,si,s Dode China North America'SoLrthArncrroa .1. rrigro L. Black lalnut E.IIS: rvidclvplantcd J. citrereu L. Ilutternutor uliite rvalnut E.LISand Canad:r .L trrttjrtr (1brr.) Ileller ArrzonaualtrLrt S.WtlSi. Mcrico J. rticrocttrpo '",r'alrr. Berl. Lillle tt or Texasblack vu,altut S WUS .1. ltin.l.sii (.Ie pson) .Iepson Claroor N.Calilbrnial,alnut Calilbrnia J. c'olifonrica S. Wats. Calilbrrrialralnut Caliibrrria ,J olorcluno Shndl. & Williarns CenlralArtrcrir:an r.r,alnut Mesico.Ceulr'al Anrcrioa .J. rnolli.s Engclrn. er llenrsl Mcrico. Guatcllala ,1. pv'ifitrnti.s Liebntan Mexico J. .jorttaicensi.sC. DC Nogal or WestLrdies u,alnut West Irrdics .J. d tt.t It.dI i.\ Griseb. Argentrna.Rolivia J. ttt'otropico Dicls Lcuador.Colornbia Hvblids Juglans spp.,juglone ancl allelopathy .1. intcrnrcdio = J. nipya x J. regia J. quttdrungulota - J. cinerea x J. regio Since antiquity',the walnul tree is knou'n as a sourcc of substanccshannlul 1o other lir.ing things. For example. rotting fruits wcre usedbl' native North Atnericaus to stun fish and decoctionsof leal'eshave long been used b.vfisltertncu to drive \\'onlts to the soil surface. Many American folk uses for u'alnut. including foliagc as a repellentfor ants and flies. have been described(313). Such uas the balcful rcputatiouof the n'alnut in parts of Itall'. that it is called "s'itchcs' trec" (3()2). Holq,cr. u'ahlut also providesamongst the earliestrecorded suggcstions of allelopalhr and to this da1'u'alnut provides tlre best paradigm of allelopathv.r.r hich has attractcd nluchatlention froln the popularpress (3-1.63. 9-5.215. 251) and conrmuuitrscnicc prograrns(99. 126,138.. 139). 2. HISTORICAL BACKGROUND Earlr nritings aboul the ualnul concernthe Old World spccics.luulunsregitr- thc courmonor Persianwalnut. The earliestrecordcd statcmenl rcgitrding thc harmful effeclsof the rvalnuttrces came frorn the Rolnanauthor Varro (309). uriting in about "1.f, 36 B C.. Jbr itlstance,/re [your neighbourJho.s ott ook-sntvc ort llrc (()nu)ton hounrlor.t,,rou vould be wrong to planl olive lree,sott tha edgc rtf suclt o votttl,.fitt^ tha,setree,s have a natural atrtipath.vto il ,srtgreat lhat, not onl.v,dttlhe.r.,heor vttrlt', but even, itt their efforts to escope,bend avo.v itttt'onlstoworrl lhe /ontr preci.sel.r'u.s tloe,stlte vine i.[planlecl neor cabbage. Like oak lree.s,trolnul lrac.srreur.r'our fitrrtr. i.f of lorye size ond stdtlding at little distance.fronrone onolht:r,make il.: nrar.gin.sl<tlall.t' unltrcductit'e."Pliuy (224) furlher expandcdon this: "[Thc shadcl o/ the walnut i.s hcov.t,anderen ceuse,sheodache in rtruttl anclirtjur-t'to an.vlhitrgplonled in it,s licinit.t'." and "Zo encJtkind o.fplant,shade i.seilher a nurse or el,sea.slcp-tttolht,r - al all evenl.s.fitrthe shadowof a volnul lree or e:ilone pitte or o,\pntce or a.silvtr fir to louchott.v plont vhatever i.t uncloubtedl.vpoison." Hc also notcsthal. "'llte ook uutl the olive are partecl b.vsuch inveterote hated thal i.f ona be plonlctl itr tlte holt,.frottr rrhtch the olher hasbeen tlug out, lhe.r,tlie.lhe ook intleed ol.vt tl.r.,ingif planlctl ttt'rrr the walnLrt." These notions were courmonll'reiterated and sourclinrcsaugurenled in subsequentnatural historl' and agricultural lracls in thc Mcdicral period. Of pa(icular intercst is tlte Andalusianregiou rvhich had a rich historr of agricullural uriters beginning rvitlt Columella. The Arab rvorld had a numbcr of grcat libraries as thc Arabs.unlike tlte Europeans.ltad accessto paper:holever" thc \ast urqoritr of lhc booksuere bumt during the Christianpurges of the l-5thand l(rllr ccnlrrrics. Alabic science.maiulv under the influence of Jabir ibn Hai1"am.developed a con-rplexsJstelr of antipathiesand s1'rnpathies,based largelv on supposedcornpositions of tltings fronr tl'iilis the elements of water, air, earth and fire (l6l). The few surviving agricultural rvritings frorn this era seem to have a rnore practical basis. The writings of the l2th centur)' Andalusian rvriter Ibn Al-Arvlarn rvere largell' based on a number of predecessorsbut theservorks. notably that of the lOth celtury Persian. Ibn Waltsltii'va have printed been recently. al-Filaha al l,iabati.v.traor Nabathean Agricdlture. cornpiled by Ibn Wahshilya, may owe its origins to ancient Babylonian lritings and. thus. predate Grcek and Roman rvritings. but the earliest knou'n r,ersion in Arabic dates to the 8th century'. Ibn WahshiS'a (137) statesnumerous plant s1mpathiesand ar(ipathiesbut thesecomrnonll'retain an occultinfluence. In describingthc rralnut he sa1-sthat it is "the opposite o.f the pomegranote vith respecl t<t cold and heot. lt t.s nrosll.y'hot." In the context of the era. the referencesto "heat" likell' rclatc lo thc bitternessQuglone), as later it is stated: "it is rare for the v,alnul kt be atlackecl h.t' tlisease.rhecau.re the *ee is strcng, hard and hot and thu.scan prorecl it,self [ronr di,sea.veand u'eakness." and "It is ltarnrful becausethe heat of it.s.fruit is intens<:nncl the heat cattsespirnples anel black morks." Ibn Al-Arvlarn's rvork. u'hile importanl in his cra. rvas effectir,'el1'lostuntil the l9th centurv rvhen translatiousappeared rn Spanish(135) and French(136). althoughit rvasevidentlvknoln to the l(>thcenlur.\ u'riter, Herrera (129). Ibn Al-Awwam wrote of the n'alnut (translaledfrom l36): ".,1// treesplanted in the viciriitll dernonstrateantipathy, tritlt tlte exc:eptiono.[lhe.lig, rhich is.fountlto have severalp<tittts of agreeabilit.lt." Hadj of Grenada says "That the walnut is antipathetic to nto.i trees tllat one vishes to plant in its vicinit.v',except thefg attd the ntul.berr.t',because the volnut is of' excessivescorching and drytress,which is harmful to allwhich cotne neor it and thiclt Qre not s.vttrpatheticto il. It destro),sever.vthingv,hich grov,.t^berteatlt it, ex(:eptcertoitl winter plants, orfern.v pla,'tts,v,hich one can grov beneothir,y bronche.s y,hen it is bore of ils leaves; when one wa,tl,t to ossociate u'ith cliubing grapevine.s.tlte.t'do tr<tl succeedat all and fail at the utter limit af enfeeblemetrt." The l3th centurv rvriter Albertus Magnus (2) described thc n'alnut as unfavourable to surrounding cultivated plants becauseof lts "indvelling exlretne loxic bitternes,i'. Pietro di Crescenziin his rvork Agricctltura Cotnm<tdorurn(62). uritlen in 1304 and printed in 1471, stated that ualnuts were hannful lo surrounding trees. Cardano (.t8) attributed the harrnful effects of n'alnut to the shacle-assisted accutnrrlationof "vapours' The writings of Pliny and other Latin authorswere compiledaud augln€lltcd rvith information from lbn Al-Awrvan and other Hispano-Arabic u'ritcrs b1-Herrera (129). He recordsthat the Castiliannalne for l'alnut hasthe mearing of "to hann"- although it is norv generally regardedthat Romance langrngc uanles for u'illuut such as nogal(Spanish), no.ver(French) and noce (Italian) are derived frour the Lati[ rrr,r for nut. This etyrnologicalmisconception is paralleled in English. *'ith thc ternr ,/uglon,s.spp., luglotte and alleiopathr "roxiorrs" derived fronr the Latin noxa. meaning hann. Curiouslr. Fuchs (94) in l-550 cleirnedthat the unrelated Greek trord for u'alnut kar\.,onuas llso derircd frour a uord nteaningcausing lteadache. The first indication of controvers,vconcenling the hannful cffccts of lalnttl appcars in l(16{ in thc writings of Erelvn (86). Evell'u r\;ts a grcal proponenl of plauting nalnut in England and he dcscribcd hou.
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