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Behavior Genetics, Vol. 6, No. I.

1976

LETTER TO THE EDITOR


Courtship Stimuli in Drosophila melanogaster
H. C. Bennet-Clark,' Mauri e Do!,' "rthur #. E!in$,' "u%re& Mannin$' an( )lorian *on S hil her'
Received /R Nov. !97

"lthou$h !e ompletel& a$ree !ith the statement in " *erho++ an( Ri har(son's ,-./01 arti le that au(io an( *isual ues, at least those pro(u e( %& !in$ *i%rations, are not re2uire( +or D. melanogaster to mate, !e stron$l& (isa$ree !ith !hat !e take to %e the impli ation %ehin( it, i.e., that au(itor& stimulation pro*i(e( %& the males' !in$ (ispla& pla&s no part in ourtship. #e +eel on+i(ent that the authors inten(e( this impli ation to %e ma(e %e ause o+ the some!hat (ismissi*e manner !ith !hi h the& re+er to the !ork o+ Bennet-Clark an( E!in$ ,-.3/, 1968), !ho on lu(e( other!ise. #e are also at a loss to un(erstan( !h& " *erho++ an( Ri har(son +in( it ne essar& to (en& the importan e o+ a ousti stimuli !hose e4isten e in no !a& a++e ts the main on lusions o+ their o!n re sear h. "lmost e*er& stu(ent o+ animal %eha*ior re o$ni5es that i+ a response an o ur in the a%sen e o+ a postulate( stimulus this tells us onl& that the stimulus is not an essential prere2uisite o+ the response. It ertainl& (oes not allo! one to on lu(e that the stimulus is +un tionless. This kin( o+ ar $ument !as thrashe( out lon$ a$o %& Tin %er$en , 1948) !hen ans!erin$ riti isms o+ the releaser on ept. The e*i(en e !hi h " *erho++ an( Ri har(son present +or the minor role o+ !in$ (ispla& is not impressi*e. The num%er o+ +lies is tin& an( the e4perimental on(itions ar-e not a(e2uatel& (es ri%e(. )urther, the time s ale o+ the ma6orit& o+ the e4periments is 2uite unrealisti 7 D. me!ana gaster ourtship, at least +or out%re( populations, is omplete( !ithin minutes, not hours or (a&s. #e ma& %e +or$i*en i+ !e (o not onsi(er that the e*i(en e presente( %& " *erho++ an( Ri har(son out!ei$hs e*i(en e amasse( in this an( other la%oratories o*er man& &ears in*ol*in$ thousan(s o+ o%ser*ations un(er a *ariet& o+ e4perimental on(itions ,Sturte*ant,

Department o+ 8oolo$&, 9ni*ersit& E(in%ur$h, E(in%ur$h, S otlan(.


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Behavior Genetics, Vol. 6, No. I. 1976


pu%lisher.

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Bennet-Clark, Do!, E!in$, Mannin$, an( *on S hil her

-.-?@ Spieth. -.?;@ Basto k an( Mannin$. -.??@ 7E!in$. -.30@ Bennet Clark an( E!in$. -.3/@ Burnet eta! .. -./-@ Bennet-Clark eta!., -./A@ *on S hil her, -./?1. "ll o+ this !ork supports the on lusion that the !in$ (is pla& sho!n %& male Drosophila +un tions to stimulate the +emale (urin$ ourtship an( to ser*e +or spe ies i(enti+i ation. " onsi(era%le proportion o+ the stimulation must %e au(itor&, althou$h in some spe ies ,e.$., D. sim"lans, Mannin$, -.?.a@ D. s"#"$ii, Mannin$, -.3?@ D. s"%o%sc"ra, Bro!n.B -.3?@ an( some o+ the D. nas"ta su%$roup, Spieth. -.3.1 there are pro%a%l& important *isual omponents. #e are $reatl& intereste( in " *erho++ an( Ri har(son's results on the role o+ pheromones, parti ularl& as !e ha*e not in*esti$ate( this un (ou%te(l& important aspe t o+ ourtship oursel*es. #e ha*e, ho!e*er, stresse( the pro%a%le primar& importan e o+ hemi al ommuni ationC in spe ies i(enti+i ation an( in maintainin$ se4ual isolation ,Mannin$, -.?.%, -.3?@ E!in$ an( Mannin$, -.3/@ E!in$, -./>, -./?1. #e think it use+ul to attempt the (istin tion %et!een stimuli !hose +un tion is se4ual stimulation an( those !hi h +un tion +or the maintenan e o+ se4ual isolation. althou$h, o+ ourse, some stimuli !ill %e in*ol*e( in %oth. #e think that hemi al stimulation ,parti ularl& in out%re( populations1 is unlikel& to %e important in raisin$ the re epti*it& o+ D. me!anogaster +emales on e ourtship has %een initiate(. "n& h&pothesis !hi h postulates su h a role +or s ent must e4plain !h& remo*al o+ the arista ,the (istal portion o+ the antenna1 alone re(u es re epti*it& o+ +emales %& as mu h as total antenne tom& (oes ,:etit, -.?D@ Mannin$, -.3/1. The ol+a tor& sense or$ans o+ the (ipteran antenna are lo ate( on the +la$ellum ,the pro4imal portion1 ,)lu$$e, -.A01. The arista a ts as a sail an( thus mo*es the !hole antenna !hen soun( parti le *elo it& !a*es, similar to those pro(u e( %& the males' !in$ (is pla&, rea h the +emale's hea(. "t the lose (istan e +rom !hi h the male ourts, an( in this parti ular a ousti re$ime, the soun( parti le *elo it& is likel& to e4 ee( D> (% an( thus is per eptuall& *er& lou( ,Bennet-Clark, -./-, -./?1. If s ent stimuli in rease the re epti*it& o+ +emales, .it is also har( to e4plain !h& !in$ remo*al +rom males $reatl& re(u es matin$ spee( ,al thou$h not ne essaril& ultimate matin$ su ess1. Certainl& this re(u tion in spee( annot %e attri%ute( to (ama$e, as, in ontrast to "*erho++ an( Ri har(son, our mortalit& rate +rom !in$ amputation is onsi(era%l& less than IE. Rein+or in$ this *ie! is the +a t that a su%stantial portion o+ male su ess an %e restore( %& arti+i iall& pro*i(in$ the appropriate a ousti stimulus ,Bennet-Clark an( E!in$, -.3/, -.3.1. Both a ousti an( hemi al stimuli ha*e %een sho!n to %e in*ol*e( in the ourtship o+ man& Diptera ,E!in$, -./?1 an( it shoul( o asion no sur prise that the same is true also o+ Drosophila. Rather than (eni$ratin$ the role o+ an& one stimulus, !e shoul( attempt to e*aluate all the omple4 o+

.S
Courtship Stimuli in Drosophila melanogaster

intera tin$ +a tors in*ol*e( in Drosophila ourtship as !e no! kno! enou$h to reali5e that an& simplisti e4planation !ill %e ina(e2uate. REFERENCES
"*erho++, #. #., an( Ri har(son, R. H. ,-./01. :heromomil ontrol o+ matin$ pat erns in Drosophila melanogaster. Behav. Genet. 07;>/-;;?. Basto k, M., an( Mannin$, ". , -.??1. The ourtship o+ Drosophila melanogaster. Behavio"r D7D?----. Bennet-Clark, H. C. ( -./-1. " ousti s o+ inse t son$. Nat"re ;A07;??-;?.. Bennet-Clark, H. C. (-./?1. Soun( pro(u tion in inse ts. &ci. 'rogr. ()*+ord, 3;7;3A-;DA. Bennet-Clark, H. C., an( E!in$, ". #. ,-.3/1. Stimuli pro*i(e( %& ourtship o+ male Drosophila melanogaster. Nat"re ;-?733.-3/-. Bennet-Clark, H. C., an( E!in$. ". #. ,-.3.1. :ulse inter*al as a riti al parameter in the ourtship son$ o+ Drosophila melanogaster. -nim. Behav. -/7/??-/?.. Bennet-Clark, H. C .. E!in$, ". # .. an( Mannin$, ". ,-./AF. The persisten e o+ ourtship stimulation in Drosophila melanogaster. Behav. Bioi. D7/3A /3.. Bro!n, R. G. B. ,-.3?1. Courtship %eha*iour in the Drosophila o%sc"ra $roup. :art II. Com parati*e stu(ies. Behavio"r ;?7;D- A;A. Burnet, B., Connoll&. K .. an( Dennis, L. ( -./-F. The +un tion an( pro essin$ o+ au(itor& in +ormation in the ourtship %eha*iour o+ Drosophila melanogaster. -nim. Behav. -.70>.-0-?. E!in$, ". #. , -.301. The in+luen e o+ !in$ area on the ourtship %eha*iour o+ Drosophila melanogaster. - nim. Behav. -;7A-3--A;>. E!in$, ". #. ,-.3.1. The $eneti %asis o+ soun( pro(u tion in Drosophila pse"doo%sc"ra an( D. persimilis. -nim. Behav. -/7???-?3>. E!in$, ". #. ,-./>1. The e*olution o+ ourtship son$s in Drosophila. Rev. .amp. -nimal E!in$, ". #. ,-./?1. Communi ation in Diptera. In Se%eok. T. ,e(.1, /o0 -nimals .omm"ni cate, In(iana 9ni*ersit& :ress, Bloomin$ton. E!in$, ". #., an( Bennet-Clark, H. C. ,-.3D1. The ourtship son$s o+ Drosophila. Behavio"r A-7;D/-A>-. E!in$, ". #., an( Mannin$, ". ,-.3/1. The e*olution an( $eneti s o+ inse t %eha*iour. -nn. Rev. 1ntomol. -;7---?- ---D. )lu$$e, C. ,-.A01. Geru hli he Raumorientierun$ *on Drosophila melanogaster. 2. Verg!. 'h3siol. ;>703A ?>>. Mannin$, ". ,-.?.aF. The se4ual %eha*iour o+ t!o si%lin$ Drosophila spe ies. Behavio"r -?7-;A-0?. Mannin$, ". ,-.?.%F. The se4ual isolation %et!een Drosophila melanogaster an( Drosophila sim"lans. -nim. Behav. /73>-3?. Mannin$, " ..,-.3?1. Drosophila an( the e*olution o+ %eha*iour. Vie0points in Biolog3 07-;?-3.. Mannin$, ". ,-.3/1. "ntennae an( se4ual re epti*it& in Drosophila melanogaster +emales. &cience -?D7-A3--A/. :etit, C. , -.?D1. Le (eterminisme $,meti2ue et ps& ho-ph&siolo$i2ue (e Ia ompetition se4uelle he5 Drosophila melanogaster. B"ll. Bioi. .;7;0D-A;.. Spieth, H. T. ,-.?;1. Matin$ %eha*ior !ithin the $enus Drosophila ,DipteraF. B"ll. -m. 4"s. Nat. /ist. ..7A.. 0/0. Spieth, H. T. , -.3.1. Courtship an( matin$ %eha*ior o+ the Drosophila nas"ta su%$roup o+ spe ies. 5niv. 6e*as '"%!. 3.-D7;?? ;/>. C Sturte*ant, ". H. ,-.-?1. E4periments on se4 re o$nition an( the pro%lem o+ se4ual sele tion in Drosophila. J. -nim. Behav. ?7A?- A33. Tin%er$en, <. , -.0D1. So ial releasers an( the e4perimental metho( re2uire( +or their stu(&. 7ilson B1.11l. 3>73 ?-. *on S hil h r, ). , -./?1. The role o+ au(itor& stimuli in the ourtship o+ Drosophila melano gaster. -nim. Bt8hav. ,in press1.

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