Donald & Nordlund 1976

7/30/2019 Donald & Nordlund 1976

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J. C!lml. &01., 1976, Vol. 2, No. 2., PIl. 211-220

TERMINOLOGY OF CHEMICAL RELEASING STIMULIIN INTRASPECIFIC AND INTERSPECIFIC

INTERACTIONS 1 20

DONALD A. NORDLUND and W.J. LEWISSouthern Gran lnsects Research Laboratoryo Agrlcultural Research Savlce

USDA, Tifton, Georgla 31794(Recelved August JI 1975; revlsed Octaba 27, 1975)

Abslracl-The lerminoJogy of chemicaJ rcleasing slimuli is exarninedin an allernpl lo reduce sorne apparenl confusion. Two new c\asses ofinlerspecific chernical signals, synornone and apneurnone, are proposed.Key Wonls--chemical ecology, hormone, pheromone, allomone, kairomone, synornone, apneurnone, chemical cornrnunicalion.

INTRODUcnON

Numerous chemicals are produced or acquired by plants and animals thatserve as releasing stimuli within or between organisms. The study oC thesechemicals and the interactions they mediate is a rapdly growing field withinthe relatively ne"'; area of chemical ecology (Sondheimer and Simeone, 1970;Whittaker and Feeny, 1971).Since the terminology describing these re1easing stimu Ji has been criticiz.ed and perhaps misunderstood, we have examined it with the hope oCeffecling sorne c1arification. Ratiler lhan present an cxhauslive list aod discussioo oC tbe chemicals and the nteractioos they mediate, we will disctJssonly sorne of the more interesting and pertinent examples. Also, we havelimited the discussion of hormones and pheromones because they ha ve beenthorougbly reviewed previously by others and appear lo be well understood.1 In cooperaljon wilh Ihe Universily of Ocorgia College of Agricullural ExpcrimenlSlalions, Coaslal Plain SIal ion, Tifton 31794. Received for pub Jication Augu sl 11, 1975.2 Mention of a proprielary product in this paper does nol constilule c:ndorsemenl of lhisproducl by Ihe USDA.

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21312 NORDLUNO ANO LEWl9Chemical agenls are c1assified acconJing lo their fundion or effecl in

specific inleraclions (Whittaker nnd Feeny, (971), and these fundions areno' mulually exclusive (Orown et al., (970). Jl is no l al all uncommon fora chemicallo fundon in tw or lhree diITcrent types of nteradions.

We do not believe that dcfinilions of"communicalion" should nffect lhevalidity of the lerminology useu for lbese releasing stimuli. For cxample,acconJing to lhe dcfinilion of BurghanJt (1970), chemicnl agcnts c1assifiedas pheromones involve lrue cOlllmllnication, whcrcas kairornones and manyallomoncs would no1. However, we do not feellhat lhe c1assincalion implieslhal the terms are eqllated comnlunicatively, a concern expressejd by Blum(1974). lmleed, as noted by Otte (1974), some of the releasing stimuli discussed are signals that are fashioned or maintaioed by natural seleclionbecause they convey information related to survival, while other sjgnals serveas incidenlal cues or signs in a parlicular relalonship but their effect in lhisrelationship is not their raison d'etre.

HORMONES

Hormones are one of the lwo major types of chernicals that mediatebehavior or physiological inleractions; semiochemicals are lhe olher. Hor-mones are chemical agents proJuced by tssues or enJocrine glands that causespecific reaclions within the producing organismo Important processes inplants and animals that are controlled by hormones inelude: phototropismand geolropism in plants, Jiapause, growth and development, digestion,sexual maturation, oestrous and menstrual cycles, lactation, and pregnancy.)formones are generally c1assiIied on the basis ofthe process they control suchas growth hormone, molling hormone, an d sex hormone. V,ft.rious aspectsof hormones have been revieweJ by Gorbman (1959), Gorbman and Bern(1962), Pincus anJ Thirnann (1948-64), Young (1961), Novak (1966), Gabe(1966), van Je r Kloot (1960), Andus (1959), Meyer et al. (1960), Siddal(1970), Engelmann (1968), and Williams (1970).

SEMIOCHEMICALSThe term semiochemical (Ok. simeon, a mark or signal) was proposed in

1971 by Law ane! Regllicr to describe lhe chemicals involved in the chemicalinteraclions between individual organisms. The semiochemicals are sub-divided into lwo major groups, pheromooes and allelochernics, dependiog onwhelher the interaction s ntraspecHic or interspedJic, respectively... P/eromolles. Pheromones have "een known to exist since at least J609

nRMINOLOOY Ol' CIIEM1CAt RELEASINO STlMULI

wheo Charles Butler described bow lone bees are atlracted aod provokecl lomass stiog by a substance re1eased by a single sling. But l was no l until 1961that tbe first isolatlon snd identification of a pheromone were reported(Outcoandl el al., 1961), and lhe sting pheromone of the honey bee, Apisml/lfera L., was no t identified unlil1962 (Boch clal., 1962). Since 1961, withthe improvements in chemical technology, literally hunureds of pheromones,

L< componenls of pheromones, and other semiochemicals have been isolatedaod identified. Pheromones have beeo demonslrated jn organisms rangingfrom slgae (Starr. 1968; Siegel and Cohen, 1962; Mucller et aL, 1971)to primates (Michael et al., 1971; Curts et a!., 1971), an J Uley have beenposlulated in humans (Comfort, 1971; Michael et al., 1974).

Tbe terro ectohormone was proposed in 1932 by Beth lo describe thechemicals involved in intraspecific interaclions, bu t the term was a cootradiction in itself (Karlson and Dutenandt, 1959) so jt was replaced by the termpheromooe (Gk. phereum, to carry, and lIOrmon, to excite or to stimulale,Karlson aod Buteoandt, 1959; Karlson and Luscher, 1959). Pheromone wasoriginally defined as a substance secreted by an animal to the outside thatcauses a specific reaction in another e m ~ r or members of the same species.Sorne more cornmon types a r e l t r a i l ; " " m ~ ~ k l l l g or trail-following pheromones,alnrro pberomones, dispersants, territoriality pberomones, synchronizationpheromones, species aggregati on pheromones, and sex pheromones (Wilsol,1963, 1965, 1970; Wilson an d Bossert, 1963; Beroza, 1970; Jacobson andBeroza, 1964; Jacobson, 1965, 1966; Butler, 1964, 1967, 1970; Regniet IlndLaw, 1968; Blurn, 1969; B1um and Brandt, 1972). A single pherornone mayhave more than one pheromonal funclion or efTe t (Morse, 1972). Perceptionof a pheromone may result in an immediate behavioral response (releasereffect) or a complex set of physiological responses that are simply sel inmotion by the inilial perception (primer elfect) (Wilson an d Bossert, 1963).These effects are also applicable to allelochemics and will be discussed Jater.

A/lel!?chemics. The lerm allelochemic was proposed by WhiUaker(1970a, b) to describe the chemicals involvcd in interspecific interactions.It was originally defined as a chemical significanl to organisms of a speciesdilferen! from its source, for rensons other than food as such. The lermllenomone was proposed by Cherniu (1970) and has essentially the samemeanlng. The term allclochemic appears to be in more general use and willbe used bere. Allelochemics are herein divided into four subgroups: allomones, kairoOlOnes, synornones, and apneumones; lhe division is based 00whether the emitter, the receiver, or both benefit in the interaction.

An al/omone is an allclochcmic originally defined as n chemical substaocc, produced or acquired by an organism, which, when it contacts anindividual of al10ther species in lhe natural context, evokes in Iltc receivera behavioral or physiological reacUon adaplively favorable to the emitter

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214 NOJtomU ANO lilwliI

(Drown, 1968). Tbe term was derived from alloiohormone, which bas essentially tbe same meaning and was proposed by Beth (1932). Allomones neludea wide variety f substnnces ranging from venorns and repellent seereUotlsin animals and plants to growth inhibitors such as allelopatbic substances andsorne nntibiotics. As with pheromones, interactions mediated by allornonesmay involve a complex sel of physiological changes (primer eITect) or animrnediale behavioral response (releaser effect). Sorne important reviews ofallomones and their effects are those of Whiltaker (l970a), Went (1910),De.lhier (1970), Williarns (1970), Eisner (1970), Whiltaker and Feeny (1911),Happ (1913), and Bucherl et al. (1968-71).Another type of allclochemic is theJt.airomone. The tetm kairomone (Gk.kairos, opportunislic) was proposed by Brown el al. (1910) and was originallydefined as a chemical subslnnce, produced or acquired by an organism, which,when it contacts an individual or another species in the nntural conlext,evokes in the receiver a behavioral or physiological reaelion adaptivilyfavorable to the receiver but not to the emitter. Thus, kairomones ate maladaplive or al leasl nonadaptive to the individual emilter, but, as Browh et aJ.(1970) pointed out, it is possible for a kairomone lo be adaptively favorableto the population as a whole by helping tegulale the populatiott dynamicsin a way lhat is favorable in the long termo For exatnp!e, Gilbert (1966)presents data showing thal a predaceous rotifer produces a substance thatcauses the unclea ved eggs of its ptey to produce spined individuals which tiredifficult for the predator to eal. This kairotnone ae!s to maintain tbe population of the predator al a level that will not destroy its food source. , [\".1'>These chernicals may be incidenlal cornpounds utiliied as CUes by lhereceiver or signal emissions ntended as pheromones, altomones, ot botmonesfor lhe legitimate receiver, bul which are exploiled by illegWmate receivers.For these reasons there has been considerable controversy about this termoBlurn (1974) argues that "the so-called,j kairomoneS appear to be pheromones nnd allomones that have "evolutionltrily back-flred'; I1nd such donot "represent a c1ass of chemical signals distinet from al1omones andpherornones." Because each specific intera dion must be consldeted seplttatelyaod these interactions are hol mutally exclusive, \Ve do hot concUr with thisreasoning. The rae! that some phagostimulants, which stimulate a Iimitednllmber ofherbivores to feed on n plenl also fundion, and probably otlginsllyevolved, to deter a vasl rnultilude of herbivorous species from feeding on thesame plant (Frankel, 1959) does not prevent lhe term kairomone froro servingas rncaningful terminology. In the lnleractions in which compound isa phagostimlllnnt, it is a kairomone and in interactions in which it is a feedingdelerrenl, it is nn allornone. Even l1 hormone can acl as a knitornone as In thecase of cortlcosleroids of lhe rabbit and the rabbit Oea, Spilopsyllw cunicull(Dale) (Rothschild, 1965).

'TAllMtNOloov o" CIlEMlcAl RElEAS1NO ~ J M U l l iB1l } ~ , '7l.' ,;,. Jo tlrder to temove sorne of tlte ambiguity preseot In sorne orthe ptevlous

b l a s s i f l c a t i o l 1 ~ , we would Iike lo propose an additional division of aUelocltemicS tor muhlalistic intetactions lhal have generally been regatdedliS.' I1lLtl1ohes (Brown et al., 1970) oc as either allomones or kairott1ohes(Whlltaker and Feeny, 1971). SynolOne (Gk. syn, witft or joinlly) is Iterdefined as acbemicat substance produccd or acquired by an organism, which,wben It contacts an individual of anolher species, in the nntural context,evokes in lhe receiver a behavioral or t>hysiological response adaptively

favorable lo both lhe emitler and the receiver. This group of allelochemicswould ioelude floral sccnts and neclars lhat altrae! insects and other pol-Iinators snd substanccs that play an importanl but ofien subtle role in! y m b i o l ~ c relalionships (llenry, 1966; Nutman and Mosse, 1963; and others).For eumple, to survive on a diet of wood, the wood-eatiog cockroach,Cryptocercus punctulatus Scudder, requires wood-digesting protozoa in itsgUl. The hormone ecdysone, which regulates molling in the cockroach, also" aets as a synomone thal induces tb.e sexual cyele of sorne of these protozoa(Clevelatld; t959), allowing them to reproduce. This is an example ot substaltce functiortiltg as a horrnohe and S synomone with a primer effeet.

Mu/tip/e Species interacilons. Clarificalion of lerminology is also b e ~ d e d for intetctiorts involving three or tnote s p e c i e ~ . interactions ihvbl-vlng tihost plnt; phytophagous insee! feeding on the plant, ltd its iIiseet t'tltaslloid. As prtviously discussed, phagostimulants that aUtad and stlmUlte thephytophagous species I1re kairomones. It is also knowh that sotne parasitoldsutilize chemical llIes emitled by the host's food plant to 10cate the hosl'shabitat (Arthur, 1962; Monteith, 1958; Ullyett, 1953; and olhers) even In theabsence of the host. We considet these cues to be synomones in this intetl1ction, sincc they benefit the receiving organism in locating ils host, and theybenefit the emitter by rl'ducing the damage caused by the phytophagous. insecto The chemical serving as a kairomone for the phytophagous speciesmay also serve as the synomone utilized by the parasitoid. Slimuli releasedfrom the phytophgous species tbat act as cues ror the parasitoid are c1earlykairomones. Vinson (1975) shows that theparasiloid Cardiocftiles nigricepsViereck js attracted to tobacco plants that have beeo damaged and that theyate stimulated into an intensive search of the surrounding planl fissue if thedamage was caused by the feeding of lIeliotlris virescens (F.). The properc!assilication of these cues is not clear. We suggest thal those chemicalsoriginating ftom tbe plant only as a resull of the aclivities oC or only incombination wilh substances ftom, the phytophagous species that Ref ascues to tbe parasitoid should be considered kairomones. These same considerations cal1 be appJied lo other multispecies interactions.There nre also three-wny interaclions in which an organism, stlcb asa parllsite oc predator, is atttaeted lo nonlivillg substances in wblch it tnll)'


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216 NORDLUND AND LIlWlsfind nnolher organism, ils host or prey, by chemical cues released frolTt thenonliving substance, even in tbe absenco of the bost or prey, Thotpe andJones (1937), for example, fouJ;ld lbat tbe ichneumonld parasite ~ e n t u r f a canescells (Oravenharst) is attracted lo the odor of its hosls' food, 1.e., freshoalmea!. Also, Laing (1937) demonstraled lbat lhe braconid Alysia monduca

" for Paozer ans! lhe chalcid NasOl/ja (= Mormonfella) vlfrlpennis (WaJker) areallracted lQ.,meal e\en Ihough it never conlained their dipleran hosts. Thesechemicals cannol be classified according lo the previously described systemfor c1assifying chemical agenls in three-way interaclions. We here wish lopropose the lerm apneumone (Ok. a-pneum, breathless or Iifeless), which wedefine as a subslnnce emitted by a nooliving material that evokes a behavioralor physiological reaclion adaplively favorable to a receiving organjsm, bu tdeltimenlal lo an organism, of anolher species, Ihal may be fouod in or onIhe nonliving malerial. Chemicals released from Ihe non living material onlyas a resull of activities of Ihe polenlial host or prey, or onJy in conjunctionwith chemicals from the potential hosl or prey thal are adaptively favotablelo the receiver, would slill be c1assified as kairomones.TABLE 1. DEfINITIONS Of OIEMICAL MEDIATORS IMPoRTN IN CHEMlcAL INTER

CTIONS MAoNG ORONISMS

A. Hormone-A chcmical agent, produced by Iissue or endocrlne (Iands, I h a ~ colllIolsvarious physiological processes within an organlsm.B. Semiochemical-A chemical nvolved in Ihe chemical J n l e r a c l i ~ belween organisms.

l. h e r o m o n ~ S u b s t a n c e Ihal Is secreled by an animal or playt lo Ihe oulslde Ihatcause a specific readion In a recelvlng IndivIdual of Ihe..same s p e c i ~ . 2. A1lelochemic-Chelnical significanl lo orgnlsms of a species dilrereilt froltt the'source, for reasons other than food as sUch.a. A l l o m o n ~ A substance, produced or acqulred by ao organistn, whlcl1, when iIcontads an individual of anolhet species In the natural conlext, evokes In thereeever a behavioral or physiological reactioo adaptlvely favorable lo lheemilter bul not lo lhe receiver.

b. K a ; r o m o n ~ A substance, produced, acquired by, or released a; D tesull ot theactivilies of an organism, which, when JI cootacls sn individuaJ of anothet speclesin lhe natural conte,;t, evokes In the teceiver a behavloral or physiologlcaJ reaclion adaptvely favorable to Ihe receiver but not to /he emitter.

e, Syoomono--A substance produced or acqulred by an organlsm whlch, when ftcontad! sn individual of another specics In Ihe oatural conlext, evokes In /hereeeiver a behavioral or physiological reaelioD ndaptlvely ravorable lo bothemiller and receiver.d. A p o e u m o n ~ A substance emilled by n nonHving material that evokes a be-haviornl or physiological reaelion adnplively favorable lo s recelvln8 organlsm,but delrimentallo no organlsm. of anolher tpecies, whlch moy be ~ o u o d In or onthe nonlivjng malerial. .

-, . ':,.':I . j' \1nMtN01.OOY 01' cllEMlcAL kEU:ASINO mMul.l 211'J 1Det6 may be olher chemical-me diated interaclions involving lon\lvJl1g)itterln! and vrious otber organisms, bu t further shJdy is heeded before tbeylta\b be c1a!lsied. At the present time, ror tlJe sake of simplicity, we are col1

. ; ~ \ " ilderlllg apnelJmones as a subgroup ofallelochemics; bu t ifmore inlerclionsr. l involving nonliving malerials are found, jt may be beneficial lo reclassifythero undet aoolher major group.

.t\CONCLUSIONSA c1assificatioo of chemical mediators that serve as re/easing slimulils ptesented in Table t. This c1assificalion is based 00 Ibe function or effeclofthe chemical in each specific inleraclio n. These functions are 00 1 mutuallyeltclusive; lihd li chemical may have a function in each of two or three difteretll types oC ioleraetions. Sorne of the original defioitions have beeo altered

~ l i g h t 1 y to accornmodale inleniclions no t previously considered and loetirninale lhe atnbiguity of sorne lerms.Th e shdy ot the chemical agents has great potential no l only troin lhe.cdemic standpoint of increasing ou t IJnderslanding of behaviot nd evoluUori, bu t also trom ptactical standpoinl, e.g., fo t controlling lhe behaviot' otvatious anitnals to ou t advanlage. Much work has been done along tms I i t 1 < ~ parlicularly wilh pherotnotles (see previous references) an d kairomoJes(Lewis et aL; t975).REFERENCES

ANO\)S, LJ. 1959. P1ant Gro lvlit Subslnnces, 2nd ed. Leonard Hill, London.ARnluR, A.p. 1962. Innuence oC hosl Iree on abundance of [Iop!eel(s eonquisilor (Say)(Hymeoopleta: Ichneumonidac), 11 polyphagous parasite of lhe European pine shooltnolh, Rhyadonia buo!fana (Schirr.) (Lepidoptera: Oleuthreutldae) Can. Enlomol.94J37-47.DERozA, M. (ed.). J970. Clrenriea!s Conlro/l(n g bueet Behavior. Academic Press, New York.DeTlI, A. 1932. Vemachfasslgle Honnone. Nafllrll'isserueha[ten 20:177-83.BLt.JM, M.S. 1969. Alarm pheromones. Annu. Rev. Entamar. 14:57-80.BLUM, M.S. J974. Dedphering Ihe eommunicalive rose Ita slone. BI/([. Entamo!. Soco 14m.20: 3(}-35.BWM, M.S. nnd BRANOT. 1.M. 1972. Socia! Inseel pheromones, their chemlstry snd funelion. 14m. Zool. 12:553-76.DoCII, R., SIII!AJU!R, D.A., ond SroNE, RC . 1962. Idenlificali on of iso-amyl scelale as aoactive compound lo lhe slng pheromone In Ihe honey bec. Nature (LoMon) 19S:JOI8-20.BROWI'I, W.l., lR. 1968. An hypolhesis conceming Ihe Cunclion of lhe melapleural g l l l l 1 d ~

In ools. 14m. Nat. 102:188-91.DROWN, W.L., lR., ErsNER. T., llnd WllnTAJt!R, R.B. 1970. AI/om ones and kalrmohM!Transpeclflc chemlcal messellgen, BloSclenee 20:21-22.


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218 219OROLtNO ANo lIl.,tmDUCIIERl, W., llucKlEv, E., and DI!ULOF1!U, V. (M.). 1968-71. Jl'tnOmolU Anlnto1J ant!

71rer Vtnornr. 3 Vols. Academlc Press, New York.DURCJIIARDT, G.M. 1970. Defining "commun(catlon," pp. 5-18. In Johnslon, J.W., Jr.,Moullon, 0.0., and Turk A. (eds.), AvanctJ In Chtmortuptfon, Vol. 1: Commun/corion b)' Chtmlcal Slgna/. Applelon-Cenlury-Crofls, New York.DUTENANDT, A., DECKMANN, R., Rnd HECItER, El. J961. Veber den SexuaJlocblofT desSeidensplnners. Tiell. Der Blologlsche Test and de hollerung des reinen Sellualloclt.slofTs Dombykol. lIoppe St)'/er's Z. Ph)'slol. Chern. 324 :71-83,DUTUR, C. 1609. 7e leminlnt monarchle. On a rrearis corrcernfng bus, an Ihl! due orl!rlng01 rhenr. Joseph Barnes. Oxford. Ciled In Wilson, E.O. 1971. 71! Instct SOcdll!s,pp. 23.5-6. Belknap. (Harvard Unlv.) Press, Cambridge, Mass.BUTlER, C.O. 1964. Pheromones in sexual processes In Inseels, pp. 66-71, In Higham, K.C.(ed.), lnstcls Rrprodllcl/on. Royal Enlomo!. SOCo (London) Symposlum No. 2.BUTlER, C.O. 1967. Inseel Pheromones. Bio/. Rev. Cambridge Phi/o SOCo 42:42-87.BUTlP.R, C.O. 1970. Chemical communicntlon In insee!s, behavioraland ecological aspecls.Advan. Chl'/floreceplon 1:3.5-78.CIlERNIN, E. 1970. Inlerspedfic chemical slgnals. BloScll!nce 20: 84S.

C l ~ V E l A N D , L.R. 1959. Sex induced wilh ecdysone. Proc. Na/. Acad. Se/. USA 4S: 741-33COMFORT, A. 1971. Likelihood of human pheromones. Na/url! (Landon) 2jO:432-33.CURns, R.F., BAlLNTINE, J.A., KEVERNE. E.B., BoNSAL, R.W., and MICHAEL, R.P. 197LIdenlificalion of primale sellua! pheromones and the properlles of synlhetic allradanls.Na/url! (Landon) 233 :396-98.DETIIIER, V.G. 1970. Chemical lnleraelions between plants and aseets, pp. 83-102, lriSondheimer, E. and Simeone, 1.B. (eds.), Chemica/ frotogy. cademlc Press, NewYork. .EISNER, T. 1970. Chemical defenses ngainst predation in arlhropods, pp. 1.57-iI8, IrISondheimer, E. and Simeone, J.B. (eds.), Chemica/ Ecotogy. cademic Press. NewYork.ENGLEMANN, F. 1968. Endocrine control of reproducllorl In Inseels. Anrlu. Rev. En/omo/.13:1-27.FRANKEl. G. 1959. The ralson d'elre of secondary planl subslances. Se/ence 129:1466-70.OAlIe, l . 1966. Nl'lIrosections. Pergamorl Press, London.Ollnl!RT, J.J. 1966. Rotifer ecology and embtyoiogical ihdudloh. Sclcnce 1.51 :12j4-37.OOIlBMAN, A. (ed.) 1959. Comparative Endocrln%g)'. Wiley, New York.OORDMAN, A. and BI!RN, A. 1962. A Tex/book 01 Comparativl! Endocrlnolog)'. Wley, NewYork.IIAPP, O.M. 1973. chemical signals belween animals. llomones and Phetomones, pp. 14990,In lIapp, O.M., Lobue, J., and Gordon, A.S. (eds.),llumora/ Con/rol olGrowrh and. DjJerenria/lon. Vol. 2: NOllverlebra/e Neuroendocrina/al)' aJid CrY/lIg. Academlc Press,New York.III!NRv, S.M. (ed.). 1966. S)'mblosls, Vol. 1. Academic Press, New York.JACODSON, M. 1965. lrruc( Su AI/rac/anrs. Wiley, New York..JAconsoN, M. 1966. Nalural nseel altraclanls and repellenls, new lools In pest con1ro/,pp. 17-26, /n Crosby, 0.0. (ed.), Na/urall'e.sl Control AgenlJ. Advan. Cheni. Ser. sj,Amer. Chem. Soc., Washinglon, D.C.JACODSON, M. and DERozA, M. 1964. lnseel allraclnnls. ScI. Am. 2JJ :20-2 .KARLSON, P. and BUTI:NANDT, A. 19.59. Pheromones (Ectohormones) la Insects. Annu. llev.En/oll/O/. 4:39-58.KARLSON, r. and LUSCIIER, M. 1959. "Pheromones" a new lerm for a c1ass of bt%glcallyaclive substances. Na/llrt (London) 183: 1.5.5-.56.

0,1. 1937. Host-finding by nseel pnrasites. 1. ObservRtions on Ihe ftnding of hosU byi' AI)'s/o manducalor, Mormon/l!lIa liftrlpennis and Trlchogramma evanescens. J. A"/trl.;\f' . &01. 6:298-3/7.LAW. 1.1-1. and RroNIElt, F.E. 1971. pheromones, Antlu. Rev. Dlochelll. 40:533-48.,: l!wlS, W.J., JONES, R.L., GROSS, 1I.R., Jr., and NORDlUND, D.A. 197.5. The role ofkalto-\. mones and olher behavioral chemicals In hosl finding by parasilic Inseels. BI!/rav. Biot.I 16:267-289.,J. MsYI!R, B.S. ANDERSON, D.B., Dnd DOIJMINO, R.ll. 1960. ln/roduc/{on lo Nant Ph)'slology.

i,LI; Van Noslrand. Prlncelon, N.J.~ ; ' . MICIlAEl, R.P., KJ!VERNE, E.B., and DoNSALl, R.W. 1971. Pheromones: Isolatlon of male, sel( allractanls from B female primate. ScellCl!. 1 7 2 : 9 ~ 6 . MICIlAJ!L, R.P., BoNSALL, R. W., and WARNER, P. 1974. Human vaginal secrelions: Volalllefatty acid conlen\. Scienu 186:1217-/9.MONTEITJI, L.G. 19.58. Influence of hosl and lis food planl on hosl-linding by Drino

'bahemlca Mesn. (Diptero: Tachinldoe) an nleruclion of olher faclors. Proc. Tenrh Inf.Congr. En/omo/. 2 :603--{)6.

MORSP., R.A. 1972. Honey bee alarm pheromone: Anolher funclion. Ann. En/omo/. SocoAn!.756:1430.,o MU!!llER; D.G.; JAENlcKe, L., DONlKE, M., and AklNToel, L. 1971. Sex aUradahl In.:! II brown algae: Chemical slrudure. Science 171 :815-17." NovAK:, vJ.A. 1966. /nSl!c/ Hormonl!s. Melhuen and CO., London.NUTroIAN, P.O. and MossE, B. (eds.). 1963. Symbio/lc Associa/ions. Cambridge UniversilyPress, Cinbridge.bTTB, b. 1974. Effecls and funclions In Ihe evolullon of signallng syslems. Annu. Rl!v. Ecot.

Sysl. S:385-417.P,Ncu.9, O. ilnd tHIMANN, KV. (eds.) 1948-64. Thl! Hormones, 4 Vols. cademic Press,New York., RllONIER, F.t. nnd LAw, 1.J-t. 1968. Inseel pheromones. J. Lipid Res. 9 :.541-.55.ROTHSCIllLD, M. 196.5. The nibbit Rea and hormones. Endmvo/lr 24 :162-68.SIDDAl, J.B. 1970. Chemical aspecls of hormonal inleraclions, pp. 281-306, In Sondheimer.E. and Slmeone, 1.B. (eds.), Clremlca/ Ecology. Acadcmic Press, New York.SII!OEl, R.W. und COIlEN, L.W. 1962. The Inlracellular difTerenlialion of cibia. Am. Zoo/.2:.5.58.STRR, R.C. 1968. Cellular differentiation in Voluox. Proc. Na/. Acad. Se/. USA 59:1082-88.SoNDIlEIMER, E. and SIMEONE. J.B. (eds.). 1970. Chtmlco/ Ec%gy. Academic Press, NewYork.TIlORPE, W.H. and JONES, F.O.W. /937. Olfaclory condilioning in R parasilic oseel and Irsrelalion lo the problem of hosl se/celion. Prac. R. Soco Landon Ser. B. 124:.56-81.UUYIlTT, O.c. 1953. Biomalhematlc and nseel populalion problems. Mtm. En/omo/. SOCoSou/Ir Alr. 2:1-89.vAN DElt KUXJT, W.G. 1960. Neurosecretlons in Inseefs. Ann/l. Rell. En/omo/. .5 :3.5-.52.VINSON, S.B. 197.5. Blochemical coevolutlon belween par:Jsltolds 8nd lheir hos ts, pp. 14-48,

'" Prlce, P. (ed.), Evo/ullonor)' S/ro/I!gles 01Parasillc lnstcts ali Milts. Plenum p ~ s , New York.W8NT, f'.W. J970. Flanls 8ud fhe chernlcal environmenl. pp. 71-82, In Sondheimer . IlttdSimeone, J.B. (ecls.), Chtmlcal frolog}'. Academic Press, New York.WIU'TTAkl!R, R.H. 19708. The blochemlcal eeology of hlgher plants, pp. 43-70, In Sdl1d.heimer, E. and Simeone, J.B. (eds.), Chtmlca/ Ec%gy. cndemic J>ress;New Yotk.WIIITTAK:IlR, RJI. 1970b. Commullfie.t and E c O l ) ' s / ~ m s . Mncmillnn Co., New York.

Jo.


6/6

220 NORDLUND AND LEWISWIIITTAUII, R.II. and FEI!NY, P.P. 1971. A J l e l o c h e m j ~ . Cherolcal fnleractlons betwcenspecies. Scimce 171 :757-70.WILLlAMS, C.M. J970. Honnonal Inleraction.5 between plants and fnsects, pp. J03-J32, inSondhemer, E. and Simeone, J.B. (eds.), Chem/ca/ Ec%gy. Academlc Press, NewYork.

WII.50N, E.O. 1963. Pheromones. Sci Am. 208:100-14.WIl.50N, E.O. 1965. Chemical communicatlons in the soclallnsects. Se/enee 149:10064-71.Wll.SON, E.O. 1970. Chemical communicatlon withln animal specfes, pp. 133-55, InSondbcimer, E. and Slmeone, J.B. (eds.), Chemiea/ Ecology. Academlc Pres.1l, NewYork.WIl.50N, E.O. lInd BossERT, W.H. 1963. ChemlcaJ communication among animals. RuenlProgr. Horm. Res. 19:673-716.YOUNO, W.C. (ed.). 1961. Sex and Inlmral Secrellofl$, (2 Vols.), 3rd ed. Wllllams andWilkins, Ballimore, Maryland.

J. Chem. Eco/., 1976, Vol. 2, No. 2, pp. 221-230

BEHAVIOR F Tefranychus urticae TOWARbESSENTIAL IL MIXTURES FRM STRAWBEltltyFOLIAGE

1.0. RODRIOUEZ,l T.R. KEMP/ and Z.T. DABROWSKt1 fJepartment 01Enlomology, Unlvers{ty 01KenllJc!cyLexlnglon, Kentucky 40506 and2 Deparlmenl 01 Ilor/icu/rure, Un{versity 01Ken/uckyLexlng/on, Ken/ucky 40506

(Recelved Augus/ 15, 1975; revised November 21,1975)

Ab!tnId- standard essential 011 mixture (SEOM) was formulatedcontaining volatile compounds In the relatlve proporlions found in theessentlal oil oflhe follage or"Citation," a strawberry cultivar relalivelyresista ni to Telranye!lUs urlleae. Olher mixtures contained varied leve/s,relative lo the SEoM, of /rans-2hexen-I-oI, nonanal, ex-terpineol, andmethyl salicylate. The behavior of T. ur/ieae femares in response lothese mixtures at several concenlralions in propylene glyco1 wasstudied in choice tube (preference) tesIs. Feeding efTecls wcre measuredby Incorporaling Ihe m i x l u r e . ~ and individual components Into sucrosewith 31p. In the choice lube tesIs, miles Were generalfy altracted bymixtures al concentralions oC 0.1 % or below unless the leve! oC metbylsaJicytate was betow 0.5 x Ihat in the SEOM, or the level oC nonanal

above Ihat oC the 5EOM. WJleh mixtures were incorporaled intofood, methyl-salicylale-stimlllaled fceding and nonanal Icvcls wereInversely relaled to lhe amollnt of Cood ingested.Key Words--strawberry resislance, TetronychlJ$ urticae, essentllll oH,.mile behavior, teeding attraclanlsfdc:lc:rrents.

lNTRODUcnONVolale organic compounds (essenlial oils) occur widely in plant t j s ~ U e s ahdare thoughllo be secondary produds of plaot melabolism. These cotnpoundsfrequenUy have very strong and characleristic odors that colllriblJle gtestlyi l>reseDI llddress: Dc:pnrtmenl of Applied Entomology, Warsaw Agricultural Unlver!ity,WllrS8wUrsynow. Poland.

221lO 1976 Plenum l'ubfllhlnl Corporlllon, 227 Ww 171h SLrl. Ne .. York, N.Y. 10011. Nd porl orlhlt publlc.Iroo 11'I0' be reproduc"d. Ilored In 1 relrle 1I)uem. or tran'rnille"'ln ony rorm or b, .ny rn ru. eloctroolc.mocbonlcal, pholocopyln,. microlllrnln,. recordln,. or o.hcrw"".. . ChOUI "Tluen permluloo or lb. publilbcr.

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Donald & Nordlund 1976