Genetics of Philaenus colour polymorphism: the 28 genotypes.

308 BRIEF REPORTS Tabk I. Numbe o b eaks in colcemid ea ed men , bo h ela i e o he HBSS ea men and cells he un ea ed con ols. The inc ease was no signi ican s a is ically. 100 cells sco ed o each concen a ion Acknowledgmm s. - This wo k has been suppo ed by Cone. To al num- wi h B eaks O gaps g an s om he Nilsson-Ehle ounda ion. My since e X 10-6 be o b eaks b eaks o pe a ec ed hanks a e due o p o . A. Le an o aluable help and mol/l and gaps gaps cell c i icism. 10 26 15 1.73 7 15 8 1.88 5 20 II 1.82 2.5 25 I1 2.27 1.4 22 I2 1.83 HBSS 19 15 1.27 Con ol 6 6 I .oo ude as ha o bu yl me cu y b omide, which was de e mined o 0.05-0.1 X lo-@ M by FAHMY (1951). The esponse o Chinese hams e cells o o ganic me cu y compounds would be expec ed o app oxima ely he same, judging om he esul s o FISKESJO (1971), who ound simila h eshold concen a ions o o he me cu y com- pounds in di e en es sys ems. The h eshold alue o HMB o c-mi osis in Chinese hams e cells was somewha highe han co esponding alues in he Allii n es . The s eng h o he e ec s dec eased wi h alling concen a ions. Fig. 1 shows ha he e was a co ela ion be ween he concen a ion o HMB and he mani es a ion o cell damage. Toxic and c-mi o ic cells we e ound mos ly in he s onges concen a ions. The ansi ions was g adual be- ween c-mi osis and no mal mi osis. The numbe o ch omosome b eaks was clea ly ele a ed bo h in he ea men s wi h HMB and wi h HBSS. This e ec was p obably due o he s a a ion he cells we e submi ed o du ing he ea men . In he a ec ed cells, howe e , he numbe o b eaks and gaps pe cell was highe in he HMB ea - Ins i u e o Gene ics, Uni e si y o Lund, Sweden Li e a u e ci ed BOYER, P. D. 1959. Sul hyd yl and disul ide g oups o enzymes. -~~ In The Enz. mes 2 ecl. (Eds. BOYER, LARDY and MYRBACK), Vol. I, Acad P ess, New Yo k, p. 51 1- 588. FAHMY, F. Y. 1951. Cy ogene ic analysis o he ac ion o o some ungicide me cu ials -- Ph. D. Thesis, Ins . Ge- ne ., Lid, Sweden. FISKESJO, G. 1969. Some esul s om Allium es s wi h o ganic me cu y halogenides. - He edi as 62: 314-322. - 1970. The e ec o wo o ganic me cu y compounds on human leukocy es in i o. - Ibid. 64: 142-146. - 1971. The e ec o wo me cu y compounds on lyso- genic E. coli K39 (k). - Ibicl. 69: 135- 138. KRISTOFFERSSON, U. 1971. The e ec o cyclama e and saccha in on he ch omosomes o a Chinese hams e cell line. lbid. 70: 271 -282. LEVAN, A. 1971. Cy ogene ic e ec s o hexyl me cu y b omide in he Allium es . - J. Indian Bo . SOC. 50A: 340 --349. RAMEL, C. 1969. Gene ic e ec s o o ganic me cu y com- pounds I. Cy ological in es iga ion on Alliuni oo s. - He edi as 61: 208 -230. RAMEL, C. and MAGNUSON, J. 1969. Gene ic e ec s o o ganic me cu y compounds 11. Ch omosome seg ega ion in D osophila melanogas e . - Ibid. 61: 231-254. RUNNSTROM, J. and MINELLI, H. 1964. Induc ion o poly- spe my by ea men o sea-u chin eggs wi h me cu ials. - Exp. Cell Res. 35: 157 STEINEGGER, E. and LEVAN, A. 1947. Cons i u ion and c-mi o ic ac i i y o iso-colchicine. - He edi as 33: 385 -396. 193. Ul K is o e sson Ins i u e o Gene ics S-223 62 Lund, Sweden 0. HALKKA, L. HALKKA, R. HOVINEN, M. RAATIKAINEN and A. VASARAINEN; Gene ics o Philaenus colou polymo phism : he 28 geno ypes (Recei ed Janua y 13, 1975) In a ecen a icle on he gene ic basis o colou ela ionships was p esen ed (HALKKA e al. 1973). polymo phism in he meadow spi lebug, Philae- The esul s o he c osses pe o med in 1972 and nus spuma i s (L.) (Homop e a), a hypo he ical 1973 we e no hen a ailable; hey co obo a e scheme o he dominance and co-dominance p ac ically all he ela ionships p oposed in he He edi as 79. 1975 BRIEF REPORTS 309 Table I. Allele combina ions and he co esponding pheno ypes in Philaenus spu na ius. The subsc ip s deno e: PT PM PL PF PC PO P PT PM i I yp PL i ma YP YP7 YP i I i4 1 i 6 ma ma ma 5 ma 5 ma la 6 yp I yp I yp P i ma la la 2 Ice lop3 K 0) expe imen al e idence lacking 1) expe imen al e idence conside ed insu icien 2) geno ype also exp essed (o belie ed o be exp essed) aspa and gib in mos indi iduals wi h app op ia e modi ie genes 3) geno ype also exp essed (o belie ed o be exp essed) as qua and alb in mos indi iduals wi h app op ia e modi ie genes 4) median s ipe o en unca ed a an e io end 5) hepa do s, o a leas he an e io do s a e p esen in mos indi iduals as inden a ions in he pale ely al ma gins 6) he lu o qua do s (see also no es 2 and 3), o a leas he an e io do s a e p esen as inden a ions in he pale 7) geno ype also exp essed (o belie ed o be exp essed) as qua ely al ma gins in some indi iduals wi h app op ia e modi ie genes He edi as 79, 1975 3 10 BRIEF REPORTS scheme. In hese yea s, 186 success ul c osses we e pe o med. They p oduced a o al p ogeny o 3799 indi iduals. Wi h se en alleles, as in Philaenus, he e a e 28 possible geno ypes (7 homozygo es and 21 he e o- zygo es). In Table 1, all he possible combina ions a e shown sepa a ely o males and emales. As explained in de ail in he p e ious a icle (HALKKA e al. 1973), he same geno ype o en p oduces dissimila pheno ypes in he wo sexes. In he able, he names o he pheno ypes a e abb e ia ed as in ou ea lie pape (see HALKKA e al. 1973 o he ull names). I no supe sc ip is a ached o he pheno ype designa ion, he geno- ype-pheno ype co espondence (g-p) has been e i ied by Mendelian F, a ios and backc osses. Supe sc ip “0” means ha he e is no i m ex- pe imen al e idence o he g-p p esen ed. In hese ins ances, he pheno ype sugges ed is hypo he ical, he e idence being ci cums an ial. O he supe sc ip s imply ha he e idence o he g-p p esen ed is based on insu icien e idence (e.g., Mendelian a ios om small p ogenies), o ha complexi ies o pene ance o exp ession exis . Some o he complexi ies a e caused by non- allelic modi ie genes (see HALKKA e al. 1973). The e ec o some o he modi ie s on pigmen a- ion is dissimila in di e en geog aphical egions, and o ms o exp ession no men ioned in he able may occu in some na u al popula ions. Such de ails do no obscu e he ac ha , wi h he help o Table 1, i is possible o calcula e allele equencies om p ac ically e e y sample collec - ed om na u al popula ions in no he n Eu ope, Asia and No h Ame ica. In Sou he n and Cen al Eu ope, many addi ional pheno ypes a e ound, bu mos o hem occu a e y low equencies (RAATIKAINEN 1971). In mos o he popula ions in es iga ed h oughou he wo ld so a he h ee homozygo es wi h uniden i ied pheno ypes (su- pe sc ip “0”) in Table 1 a e e y a e. Thei e- quencies usually a e wi hin he ange 0.005- 0.0001. In ac , all he combina ions excep p /p can be conside ed in equen in na u al popula- ions. In Denma k, Finland, No way and Sweden he equency o he allele p is o en as high as 0.9. Ve y high p equencies a e simila ly ound in o he pa s o Eu ope, and in Asia and No h Ame ica. As he sexes a eequal in numbe in newly eme ged Philaenus adul s, signi ican inaccu acy is no in oduced i only he emale sex is used in de e mina ions o allele equencies o he whole species. This is o en necessa y, because many o he alleles a e exp essed only in he emales. Ten yea s ago, HUTCHINSON (1964) w o e on Philae ius ha “- - - no clea unde s anding o he whole si ua ion, which may well p o e o be one o he mos d ama ic examples o poly- mo phism, will be possible wi hou gene ic knowl- edge - - - -”. Al hough he “gene ic knowledge” called o by HUTCHINSON will no co e he mos minu e de ails o many yea s, he in o ma ion p esen ed in Table 1 signi ies a decisi e b eak- h ough in he unde s anding o polymo phic equilib ia in he spi lebug. The many complexi- ies o gene ic de e mina ion e ealed by he able a e less de e en han hey seem. This is so be- cause in mos na u al popula ions he “clea ” geno ypes s and o well o e 90y’ o he gene pool. Acknowledgn en s - Mos o he wo k was pe o med a he Depa men o Pes In es iga ion, Ag icul u al Re- sea ch Cen e, Tikku ila, Finland. Wi hou he acili ies gene ously p o ided by he o me and p esen Heads o his Depa men , P o esso s Veikko Kane o and Ma i Ma kkula, he wo k would no ha e been possible. In a ious phases o he wo k, we ha e been aided by Si no Ho inen and Ta ja Kohila The s udy has been suppo ed by g an s om he Uni e si y o Helsinki and om he Na ional Resea ch Council o Sciences o he Academy o Finland. Depa men o Gene ics, Uni e si y o Helsinki, (M.R.) Depa men o Biology, Uni e si y o Jy askyla and (A.V.) Depa men o Pes In- es iga ion, Ag icul u al Resea ch Cen e, Tikku- ila, Finland Li e a u e ci ed HALKKA, O., HALKKA, L , RAATIKAINEN, M. and HOVINEN, R. 1973. The gene ic basis o balanced polymo phism in Philaenus (Homop e a). - He edi as 74: 69 -80. HUTCHINSON, G. E. 1964. A no e on he polymo phism o Philaenus .cpuma ius (L.) (Homop ., Ce copidae) in B i ain. -~ En omol. Mon hly Mag. 99: 175 -178 RAATIKAINEN, M. 1971. The polymo phism o Philaenus spuma ius (L.) in no he n I aly. - Ann. En omol. Fenn. 37: 72-79. Olli Halkka Depa men o Gene ics P. Rau a ieka u 13 SF-00100 Helsinki 10, Finland He edi as 79, 1975