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

211-220
TERMINOLOGY OF CHEMICAL RELEASING STIMULI
IN INTRASPECIFIC AND INTERSPECIFIC
INTERACTIONS
1 2
0
DONALD A. NORDLUND and W.J. LEWIS
Southern Gran lnsects Research Laboratoryo Agrlcultural Research Savlce
USDA, Tifton, Georgla 31794
(Recelved August JI o 1975; revlsed Octaba 27, 1975)
Abslracl-The lerminoJogy of chemicaJ rcleasing slimuli is exarnined
in an allernpl lo reduce sorne apparenl confusion. Two new c\asses of
inlerspecific chernical signals, synornone and apneurnone, are pro
posed.
Key Wonls--chemical ecology, hormone, pheromone, allomone, kairo
mone, synornone, apneurnone, chemical cornrnunicalion.
INTRODUcnON
Numerous chemicals are produced or acquired by plants and animals that
serve as releasing stimuli within or between organisms. The study oC these
chemicals and the interactions they mediate is a rapdly growing field within
the relatively ne"'; area of chemical ecology (Sondheimer and Simeone, 1970;
Whittaker and Feeny, 1971).
Since the terminology describing these re1easing stimu Ji has been criti
ciz.ed and perhaps misunderstood, we have examined it with the hope oC
effecling sorne c1arification. Ratiler lhan present an cxhauslive list aod dis
cussioo oC tbe chemicals and the nteractioos they mediate, we will disctJss
only sorne of the more interesting and pertinent examples. Also, we have
limited the discussion of hormones and pheromones because they ha ve been
thorougbly reviewed previously by others and appear lo be well understood.
1 In cooperaljon wilh Ihe Universily of Ocorgia College of Agricullural Expcrimenl
Slalions, Coaslal Plain SIal ion, Tifton 31794. Received for pubJication Augusl 11, 1975.
2 Mention of a proprielary product in this paper does nol constilule c:ndorsemenl of lhis
producl by Ihe USDA.
211
213
212 NORDLUNO ANO LEWl9
Chemical agenls are c1assified acconJing lo their fundion or effecl in
specific inleraclions (Whittaker nnd Feeny, (971), and these fundions are
no' mulually exclusive (Orown et al., (970). Jl is nol al all uncommon for
a chemicallo fundon in tw or lhree diITcrent types of nteradions.
We do not believe that dcfinilions of"communicalion" should nffect lhe
validity of the lerminology useu for lbese releasing stimuli. For cxample,
acconJing to lhe dcfinilion of BurghanJt (1970), chemicnl agcnts c1assified
as pheromones involve lrue cOlllmllnication, whcrcas kairornones and many
allomoncs would no1. However, we do not feellhat lhe c1assincalion implies
lhal the terms are eqllated comnlunicatively, a concern expressejd by Blum
(1974). lmleed, as noted by Otte (1974), some of the releasing stimuli dis
cussed are signals that are fashioned or maintaioed by natural seleclion
because they convey information related to survival, while other sjgnals serve
as incidenlal cues or signs in a parlicular relalonship but their effect in lhis
relationship is not their raison d'etre.
HORMONES
Hormones are one of the lwo major types of chernicals that mediate
behavior or physiological inleractions; semiochemicals are lhe olher. Hor
mones are chemical agents proJuced by tssues or enJocrine glands that cause
specific reaclions within the producing organismo Important processes in
plants and animals that are controlled by hormones inelude: phototropism
and 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 such
as growth hormone, molling hormone, and sex hormone. V,ft.rious aspects
of hormones have been revieweJ by Gorbman (1959), Gorbman and Bern
(1962), Pincus anJ Thirnann (1948-64), Young (1961), Novak (1966), Gabe
(1966), van Jer Kloot (1960), Andus (1959), Meyer et al. (1960), Siddal
(1970), Engelmann (1968), and Williams (1970).
SEMIOCHEMICALS
The term semiochemical (Ok. simeon, a mark or signal) was proposed in
1971 by Law ane! Regllicr to describe lhe chemicals involved in the chemical
interaclions between individual organisms. The semiochemicals are sub
divided into lwo major groups, pheromooes and allelochernics, dependiog on
whelher 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 lo
mass stiog by a substance re1eased by a single sling. But l was nol until 1961
that tbe first isolatlon snd identification of a pheromone were reported
(Outcoandl el al., 1961), and lhe sting pheromone of the honey bee, Apis
ml/lfera L., was not identified unlil1962 (Boch clal., 1962). Since 1961, with
the improvements in chemical technology, literally hunureds of pheromones,
L
<
componenls of pheromones, and other semiochemicals have been isolated
aod identified. Pheromones have beeo demonslrated jn organisms ranging
from slgae (Starr. 1968; Siegel and Cohen, 1962; Mucller et aL, 1971)
to primates (Michael et al., 1971; Curts et a!., 1971), anJ Uley have been
poslulated in humans (Comfort, 1971; Michael et al., 1974).
Tbe terro ectohormone was proposed in 1932 by Beth lo describe the
chemicals involved in intraspecific interaclions, but the term was a cootradic
tion in itself (Karlson and Dutenandt, 1959) so jt was replaced by the term
pheromooe (Gk. phereum, to carry, and lIOrmon, to excite or to stimulale,
Karlson aod Buteoandt, 1959; Karlson and Luscher, 1959). Pheromone was
originally defined as a substance secreted by an animal to the outside that
causes a specific reaction in another or members of the same species.
Sorne more cornmon types or trail-following pheromones,
alnrro pberomones, dispersants, territoriality pberomones, synchronization
pheromones, species aggregation pheromones, and sex pheromones (Wilsol,
1963, 1965, 1970; Wilson and Bossert, 1963; Beroza, 1970; Jacobson and
Beroza, 1964; Jacobson, 1965, 1966; Butler, 1964, 1967, 1970; Regniet Ilnd
Law, 1968; Blurn, 1969; B1um and Brandt, 1972). A single pherornone may
have more than one pheromonal funclion or efTe t (Morse, 1972). Perception
of a pheromone may result in an immediate behavioral response (releaser
effect) or a complex set of physiological responses that are simply sel in
motion by the inilial perception (primer elfect) (Wilson and 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 species
dilferen! from its source, for rensons other than food as such. The lerm
llenomone was proposed by Cherniu (1970) and has essentially the same
meanlng. The term allclochemic appears to be in more general use and will
be used bere. Allelochemics are herein divided into four subgroups: allo
mones, kairoOlOnes, synornones, and apneumones; lhe division is based 00
whether the emitter, the receiver, or both benefit in the interaction.
An al/omone is an allclochcmic originally defined as n chemical sub
staocc, produced or acquired by an organism, which, when it contacts an
individual of al10ther species in lhe natural context, evokes in Iltc receiver
a behavioral or physiological reacUon adaplively favorable to the emitter
.f
214 NOJtomU ANO lilwliI
(Drown, 1968). Tbe term was derived from alloiohormone, which bas essen
tially tbe same meaning and was proposed by Beth (1932). Allomones nelude
a wide variety f substnnces ranging from venorns and repellent seereUotls
in animals and plants to growth inhibitors such as allelopatbic substances and
sorne nntibiotics. As with pheromones, interactions mediated by allornones
may involve a complex sel of physiological changes (primer eITect) or an
imrnediale behavioral response (releaser effect). Sorne important reviews of
allomones 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 originally
defined 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 adaptivily
favorable to the receiver but not to the emitter. Thus, kairomones ate mal
adaplive 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 favorable
to the population as a whole by helping tegulale the populatiott dynamics
in a way lhat is favorable in the long termo For exatnp!e, Gilbert (1966)
presents data showing thal a predaceous rotifer produces a substance that
causes the unclea ved eggs of its ptey to produce spined individuals which tire
difficult for the predator to eal. This kairotnone ae!s to maintain tbe popula
tion of the predator al a level that will not destroy its food source. , [\".1'>
These chernicals may be incidenlal cornpounds utiliied as CUes by lhe
receiver or signal emissions ntended as pheromones, altomones, ot botmones
for lhe legitimate receiver, bul which are exploiled by illegWmate receivers.
For these reasons there has been considerable controversy about this termo
Blurn (1974) argues that "the so-called,j kairomoneS appear to be phero
mones nnd allomones that have "evolutionltrily back-flred'; I1nd such do
not "represent a c1ass of chemical signals distinet from al1omones and
pherornones." Because each specific interadion must be consldeted seplttately
aod these interactions are hol mutally exclusive, \Ve do hot concUr with this
reasoning. The rae! that some phagostimulants, which stimulate a Iimited
nllmber ofherbivores to feed on n plenl also fundion, and probably otlginslly
evolved, to deter a vasl rnultilude of herbivorous species from feeding on the
same plant (Frankel, 1959) does not prevent lhe term kairomone froro serving
as rncaningful terminology. In the lnleractions in which compound is
a phagostimlllnnt, it is a kairomone and in interactions in which it is a feeding
delerrenl, it is nn allornone. Even l1 hormone can acl as a knitornone as In the
case of cortlcosleroids of lhe rabbit and the rabbit Oea, Spilopsyllw cunicull
(Dale) (Rothschild, 1965).
'TAllMtNOloov o" CIlEMlcAl RElEAS1NO iB ';1

'7
l.' ,;,. Jo tlrder to temove sorne oftlte ambiguity preseot In sorne orthe ptevlous
we would Iike lo propose an additional division of aUelo
cltemicS 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 Iter
defined 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 allelochemics
would ioelude floral sccnts and neclars lhat altrae! insects and other pol
Iinators snd substanccs that play an importanl but ofien subtle role in
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 its
gUl. 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 sub
staltce functiortiltg as a horrnohe and S synomone with a primer effeet.
Mu/tip/e Species interacilons. Clarificalion of lerminology is also
for intetctiorts involving three or tnote . interactions ihvbl-vlng ti
host plnt; phytophagous insee! feeding on the plant, ltd its iIiseet t'tltasl
loid. As prtviously discussed, phagostimulants that aUtad and stlmUlte the
phytophagous species I1re kairomones. It is also knowh that sotne parasitolds
utilize chemical llIes emitled by the host's food plant to 10cate the hosl's
habitat (Arthur, 1962; Monteith, 1958; Ullyett, 1953; and olhers) even In the
absence of the host. We considet these cues to be synomones in this intetl1c
tion, sincc they benefit the receiving organism in locating ils host, and they
benefit the emitter by rl'ducing the damage caused by the phytophagous
. insecto The chemical serving as a kairomone for the phytophagous species
may also serve as the synomone utilized by the parasitoid. Slimuli released
from the phytophgous species tbat act as cues ror the parasitoid are c1early
kairomones. Vinson (1975) shows that theparasiloid Cardiocftiles nigriceps
Viereck js attracted to tobacco plants that have beeo damaged and that they
ate stimulated into an intensive search of the surrounding planl fissue if the
damage was caused by the feeding of lIeliotlris virescens (F.). The proper
c!assilication of these cues is not clear. We suggest thal those chemicals
originating ftom tbe plant only as a resull of the aclivities oC or only in
combination wilh substances ftom, the phytophagous species that Ref as
cues to tbe parasitoid should be considered kairomones. These same con
siderations cal1 be appJied lo other multispecies interactions.
There nre also three-wny interaclions in which an organism, stlcb as
a parllsite oc predator, is atttaeted lo nonlivillg substances in wblch it tnll)'
1 '
216
NORDLUND AND LIlWls
find nnolher organism, ils host or prey, by chemical cues released frolTt the
nonliving substance, even in tbe absenco of the bost or prey, Thotpe and
Jones (1937), for example, fouJ;ld lbat tbe ichneumonld parasite
canescells (Oravenharst) is attracted lo the odor of its hosls' food, 1.e., fresh
oalmea!. Also, Laing (1937) demonstraled lbat lhe braconid Alysia monduca
" for Paozer ans! lhe chalcid NasOl/ja (= Mormonfella) vlfrlpennis (WaJker) are
allracted lQ.,meal e\en Ihough it never conlained their dipleran hosts. These
chemicals cannol be classified according lo the previously described system
for c1assifying chemical agenls in three-way interaclions. We here wish lo
propose the lerm apneumone (Ok. a-pneum, breathless or Iifeless), which we
define as a subslnnce emitted by a nooliving material that evokes a behavioral
or physiological reaclion adaplively favorable to a receiving organjsm, but
deltimenlal lo an organism, of anolher species, Ihal may be fouod in or on
Ihe nonliving malerial. Chemicals released from Ihe nonliving material only
as a resull of activities of Ihe polenlial host or prey, or onJy in conjunction
with chemicals from the potential hosl or prey thal are adaptively favotable
lo 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, colllIols
various physiological processes within an organlsm.
B. Semiochemical-A chemical nvolved in Ihe chemical belween organisms.
l. Ihal Is secreled by an animal or playt lo Ihe oulslde Ihat
cause a specific readion In a recelvlng IndivIdual of Ihe..same
2. A1lelochemic-Chelnical significanl lo orgnlsms of a species dilrereilt froltt the'
source, for reasons other than food as sUch.
a. substance, produced or acqulred by ao organistn, whlcl1, when iI
contads an individual of anolhet species In the natural conlext, evokes In the
reeever a behavioral or physiological reactioo adaptlvely favorable lo lhe
emilter bul not lo lhe receiver.
b. substance, produced, acquired by, or released a; D tesull ot the
activilies of an organism, which, when JI cootacls sn individuaJ of anothet specles
in lhe natural conte,;t, evokes In the teceiver a behavloral or physiologlcaJ reac
lion adaptvely favorable to Ihe receiver but not to /he emitter.
e, Syoomono--A substance produced or acqulred by an organlsm whlch, when ft
contad! sn individual of another specics In Ihe oatural conlext, evokes In /he
reeeiver a behavioral or physiological reaelioD ndaptlvely ravorable lo both
emiller and receiver.
d. 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 In or on
the nonlivjng malerial. .
'-,. ':,.'
:I .
j'
\ \1nMtN01.OOY 01' cllEMlcAL kEU:ASINO mMul.l 211
'J 1Det6 may be olher chemical-mediated interaclions involving lon\lvJl1g
)itterln! and vrious otber organisms, but further shJdy is heeded before tbey
lta\b be c1a!lsied. At the present time, ror tlJe sake of simplicity, we are col1
ilderlllg apnelJmones as a subgroup ofallelochemics; but ifmore inlerclions
r.l
involving nonliving malerials are found, jt may be beneficial lo reclassify
thero undet aoolher major group.
.t\
CONCLUSIONS
A c1assificatioo of chemical mediators that serve as re/easing slimuli
ls ptesented in Table t. This c1assificalion is based 00 Ibe function or effecl
ofthe chemical in each specific inleraclion. These functions are 001 mutually
eltclusive; lihd li chemical may have a function in each of two or three dif
teretll types oC ioleraetions. Sorne of the original defioitions have beeo altered
to accornmodale inleniclions not previously considered and lo
etirninale lhe atnbiguity of sorne lerms.
The shdy ot the chemical agents has great potential nol only troin lhe
.cdemic standpoint of increasing out IJnderslanding of behaviot nd evolu
Uori, but also trom ptactical standpoinl, e.g., fot controlling lhe behaviot' ot
vatious anitnals to out advanlage. Much work has been done along tms
parlicularly wilh pherotnotles (see previous references) and kairomoJes
(Lewis et aL; t975).
REFERENCES
ANO\)S, LJ. 1959. P1ant Grolvlit 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 shool
tnolh, Rhyadonia buo!fana (Schirr.) (Lepidoptera: Oleuthreutldae) Can. Enlomol.
94J37-47.
DERozA, M. (ed.). J970. Clrenriea!s Conlro/l(ng 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 roseIta slone. BI/([. Entamo!. Soco 14m.
20: 3(}-35.
BWM, M.S. nnd BRANOT. 1.M. 1972. Socia! Inseel pheromones, their chemlstry snd fune
lion. 14m. Zool. 12:553-76.
DoCII, R., SIII!AJU!R, D.A., ond SroNE, RC. 1962. Idenlificalion of iso-amyl scelale as ao
active 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
In ools. 14m. Nat. 102:188-91.
DROWN, W.L., lR., ErsNER. T., llnd WllnTAJt!R, R.B. 1970. AI/omones and kalrmohM!
Transpeclflc chemlcal messellgen, BloSclenee 20:21-22.
218
219 NOROLtNO ANo lIl.,tm
DUCIIERl, 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/co
rion b)' Chtmlcal Slgna/. Applelon-Cenlury-Crofls, New York.
DUTENANDT, A., DECKMANN, R., Rnd HECItER, El. J961. Veber den SexuaJlocblofT des
Seidensplnners. 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!rlng
01 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.
L.R. 1959. Sex induced wilh ecdysone. Proc. Na/. Acad. Se/. USA 4S: 741-33
COMFORT, 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. 197L
Idenlificalion 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, lri
Sondheimer, E. and Simeone, 1.B. (eds.), Chemica/ frotogy. cademlc Press, New
York. .
EISNER, T. 1970. Chemical defenses ngainst predation in arlhropods, pp. 1.57-iI8, IrI
Sondheimer, E. and Simeone, J.B. (eds.), Chemica/ Ecotogy. cademic Press. New
York.
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, New
York.
IIAPP, O.M. 1973. chemical signals belween animals. llomones and Phetomones, pp. 149
90,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%glcally
aclive substances. Na/llrt (London) 183: 1.5.5-.56.
0,1. 1937. Host-finding by nseel pnrasites. 1. ObservRtions on Ihe ftnding of hosU by
i' 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!.
MICIlAJ!L, R.P., BoNSALL, R. W., and WARNER, P. 1974. Human vaginal secrelions: Volallle
fatty 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/. Soco
An!.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 Universily
Press, 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, New
York.
TIlORPE, W.H. and JONES, F.O. W. /937. Olfaclory condilioning in R parasilic oseel and Irs
relalion 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/. SOCo
Sou/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 hosts, pp. 14-48,
'" Prlce, P. (ed.), Evo/ullonor)' S/ro/I!gles 01 Parasillc lnstcts ali Milts. Plenum
New York.
W8NT, f'.W. J970. Flanls 8ud fhe chernlcal environmenl. pp. 71-82, In Sondheimer . Ilttd
Simeone, 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 Mncmillnn Co., New York.
Jo.
220
NORDLUND AND LEWIS
WIIITTAUII, R.II. and FEI!NY, P.P. 1971. Cherolcal fnleractlons betwcen
species. Scimce 171 :757-70.
WILLlAMS, C.M. J970. Honnonal Inleraction.5 between plants and fnsects, pp. J03-J32, in
Sondhemer, E. and Simeone, J.B. (eds.), Chem/ca/ Ec%gy. Academlc Press, New
York.
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, In
Sondbcimer, E. and Slmeone, J.B. (eds.), Chemiea/ Ecology. Academlc Pres.1l, New
York.
WIl.50N, E.O. lInd BossERT, W.H. 1963. ChemlcaJ communication among animals. Ruenl
Progr. Horm. Res. 19:673-716.
YOUNO, W.C. (ed.). 1961. Sex and Inlmral Secrellofl$, (2 Vols.), 3rd ed. Wllllams and
Wilkins, Ballimore, Maryland.
J. Chem. Eco/., 1976, Vol. 2, No. 2, pp. 221-230

BEHAVIOR F Tefranychus urticae TOWARb
ESSENTIAL IL MIXTURES FRM STRAWBEltlty
FOLIAGE

1.0. RODRIOUEZ,l T.R. KEMP/ and Z.T. DABROWSKt
j
1 fJepartment 01 Enlomology, Unlvers{ty 01 KenllJc!cy
Lexlnglon, Kentucky 40506 and
2 Deparlmenl 01 Ilor/icu/rure, Un{versity 01 Ken/ucky
Lexlng/on, Ken/ucky 40506
(Recelved Augus/ 15, 1975; revised November 21,1975)
Ab!tnId- standard essential 011 mixture (SEOM) was formulated
containing volatile compounds In the relatlve proporlions found in the
essentlal oil oflhe follage or"Citation," a strawberry cultivar relalively
resista ni to Telranye!lUs urlleae. Olher mixtures contained varied leve/s,
relative lo the SEoM, of /rans-2hexen-I-oI, nonanal, ex-terpineol, and
methyl salicylate. The behavior of T. ur/ieae femares in response lo
these mixtures at several concenlralions in propylene glyco1 was
studied in choice tube (preference) tesIs. Feeding efTecls wcre measured
by Incorporaling Ihe and individual components Into sucrose
with 31p. In the choice lube tesIs, miles Were generalfy altracted by
mixtures al concentralions oC 0.1 % or below unless the leve! oC metbyl
saJicytate was betow 0.5 x Ihat in the SEOM, or the level oC nonanal
above Ihat oC the 5EOM. WJleh mixtures were incorporaled into
food, methyl-salicylale-stimlllaled fceding and nonanal Icvcls were
Inversely relaled to lhe amollnt of Cood ingested.
Key Words--strawberry resislance, TetronychlJ$ urticae, essentllll oH,.
mile behavior, teeding attraclanlsfdc:lc:rrents.
lNTRODUcnON
Volale organic compounds (essenlial oils) occur widely in plant ahd
are thoughllo be secondary produds of plaot melabolism. These cotnpounds
frequenUy have very strong and characleristic odors that colllriblJle gtestly
i l>reseDI llddress: Dc:pnrtmenl of Applied Entomology, Warsaw Agricultural Unlver!ity,
WllrS8wUrsynow. Poland.

221
lO 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.