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Roles of juvenile hormone in the green peach aphid, myzus persicae sulzer (homoptera: aphididae) Verma, Kulbhushan 1981

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ROLES OF JUVENILE HORMONE IN THE GREEN PEACH APHID Myzus p e r s i c a e  (Sulzer)(HOMOPTERA:  APHIDIDAE)  by KULBHUSHAN[VERMA B.Sc.  (Hons.) A g r i c , H a r y a n a A g r i c u l t u r a l U n i v e r s i t y ( I n d i a ) , 1978  A THESIS SUBMITTED IN PARTIAL  FULFILMENT  OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF  SCIENCE  in FACULTY OF GRADUATE STUDIES THE DEPARTMENT OF PLANT SCIENCE  We  a c c e p t t h i s t h e s i s as c o n f o r m i n g to the required standard  THE UNIVERSITY OF BRITISH COLUMBIA January,  1981  K u l b h u s h a n Verma,  1981  In p r e s e n t i n g requirements  this thesis  British  it  freely available  for  that  Columbia,  I agree that f o r reference  permission  scholarly  f u l f i l m e n t of the  f o r an a d v a n c e d d e g r e e a t t h e U n i v e r s i t y  of  agree  in partial  the Library  shall  and s t u d y .  I  f o r extensive  p u r p o s e s may  for  that  copying or p u b l i c a t i o n  f i n a n c i a l gain  shall  Department o f The U n i v e r s i t y o f B r i t i s h 2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  Date  I o /no  \  of this  Itis thesis  n o t be a l l o w e d w i t h o u t my  permission.  Columbia  thesis  by t h e h e a d o f my  d e p a r t m e n t o r by h i s o r h e r r e p r e s e n t a t i v e s . understood  further  copying of t h i s  be g r a n t e d  make  written  ABSTRACT  The  role  of juvenile  hormone  (JH) i n a l a t e - a p t e r o u s  p o l y m o r p h i s m was  investigated  Myzus p e r s i c a e .  A t h i g h e r c o n c e n t r a t i o n s (65 ppm), t h e j u v e n i l e  hormone a n a l o g u e  i n the green  (JHA), k i n o p r e n e ,  a p t e r i f o r m nymphs.  A t lower  was  peach  aphid,  immediately  c o n c e n t r a t i o n s (10 ppm), t h e  compound was n o n - t o x i c a n d e x h i b i t e d no a p p a r e n t activity  i n apteriform stages.  administered apterizing  when t h e k i n o p r e n e the  least  I n c o n t r a s t , 65 ppm  The e x t e n t o f t h e s e e f f e c t s was  sensitive  applied.  Fourth instar  sclerotization  K i n o p r e n e - t r e a t e d t h i r d instars'underwent b e f o r e metamorphosing i n t o and p i g m e n t a t i o n  kinoprene  and d e p e n d e d upon  a l a t i f o r m s were  a s k i n o p r e n e - t r e a t e d nymphs  i n t o normal a d u l t s w i t h reduced  tization  morphological  t o a l a t i f o r m nymphs h a d j u v e n i l i z i n g  effects.  toxic to  developed and p i g m e n t a t i o n .  a supernumerary moult  a d u l t s with malformed wings. were a l s o  instar  lacking  i n these  insects.  When f i r s t  and second  alatiformsiwere treated  kinoprene,  they a l s o underwent a supernumerary moult.  a d u l t s w h i c h emerged e x h i b i t e d b o t h characteristics.  larval  and  Wing d e v e l o p m e n t was a l m o s t  lacking.  sclerotization The s e c o n d a r y  with  totally  s e n s o r i a were a l s o  inhibited;  In  and p i g m e n t a t i o n were r e d u c e d  antennal  The  apterous  t h e c a u d a a n d g e n i t a l p l a t e were p o o r l y d e v e l o p e d . addition,  Sclero-  and o c e l l  malformed.  These  findings clearly  employed  as a JH mimic  epidermal c e l l s  (1) d e m o n s t r a t e t h a t k i n o p r e n e c a n t o a l t e r the normal programming  i n a l a t i f o r m nymphs a n d  of the  (2) i n d i c a t e t h a t  p l a y s an i m p o r t a n t r o l e i n a p h i d m o r p h o g e n e s i s The d i f f e r e n t i a l  and  responses of the four a l a t i f o r m  JH  polymorphism.  nymphal  i n s t a r s s u g g e s t t h a t e l e v a t e d JH l e v e l s d u r i n g t h e f i r s t second i n s t a r s are p a r t i c u l a r l y  be  important i n i n h i b i t i n g  and wing  development. To d e t e r m i n e t h e p r e n a t a l e f f e c t s o f JH on w i n g d e v e l o p ment a n d m o r p h o g e n e s i s , newly  ecdysed  k i n o p r e n e was  apterous adults.  alate production,  also administered to  Even though c o n d i t i o n s f a v o u r e d  75% o f t h e o f f s p r i n g p r o d u c e d by k i n o p r e n e -  t r e a t e d v i r g i n o p a r a e developed i n t o normal a p t e r a e . s u g g e s t s t h a t e l e v a t e d JH t i t e r s wing development  This  i n m a t e r n a l haemolymph  and promote development  inhibit  of apterae.  Topical a p p l i c a t i o n of the a n t i - a l l a t o t r o p i n , precocene-II, h a d v a r i a b l e e f f e c t s on a p t e r i f o r m nymphs a n d a d u l t s .  Ina l l  stages, precocene produced a s i g n i f i c a n t d e c l i n e i n  larvi-  position.  and  The e f f e c t s w e r e more p r o n o u n c e d  in first  second  i n s t a r a p t e r i f o r m nymphs a n d a p t e r o u s a d u l t s t h a n i n t h i r d f o u r t h i n s t a r nymphs. insects, The  When k i n o p r e n e was  applied to these  l a r v i p o s i t i o n increased s i g n i f i c a n t l y after  2 days.  f i n d i n g s d e m o n s t r a t e t h a t JH s t i m u l a t e s r e p r o d u c t i o n i n  v i v i p a r o u s morphs o f  persicae.  and  iii  TABLE OF CONTENTS Page ABSTRACT  1  L I S T OF TABLES  v  L I S T OF FIGURES  v  ACKNOWLEDGEMENTS INTRODUCTION  i  i  x  i  1  MATERIALS AND METHODS  1  1  RESULTS  1  7  A.  1  7  1  7  2  2  2  5  2  8  B.  KINOPRENE TREATMENTS (a)  First  instar  (b)  Second  (c)  Third instar  (d)  Fourth  (e)  Apteriform  nymphs o f a p t e r o u s a d u l t s  31  (f)  Apteriform  nymphs o f a l a t e a d u l t s  32  (g)  Apterous adults  (apterae-producers)  38  (h)  Apterous adults  (alate-producers)  45  instar  instar  a l a t i f o r m nymphs a l a t i f o r m nymphs a l a t i f o r m nymphs a l a t i f o r m nymphs  COMBINED EFFECTS OF PRECOCENE AND KINOPRENE  4  5  (a)  F i r s t and s e c o n d i n s t a r apterous adults  apteriform  nymphs o f 45  (b)  T h i r d and f o u r t h i n s t a r apterous adults  apteriform  nymphs o f 47  (c)  Apterous adults  51  (d)  A p t e r o u s nymphs o f a l a t e a d u l t s  55  iv  Page DISCUSSION  61  A.  POLYMORPHISM  IN THE GREEN PEACH APHID  B.  ROLE OF JUVENILE HORMONE  61 64  (a)  Reproduction  65  (b)  M o u l t i n g and metamorphosis  68  (c)  Sclerotization  73  (d)  Cephalic sensory  (e)  Wing f o r m a t i o n  76  (f)  Lethal  80  effects  organs  74  REFERENCES  81  APPENDIX  87  L I S T OF TABLES  S e q u e n t i a l l e t h a l e f f e c t s o f 65 ppm kinoprene applied to f i r s t i n s t a r a l a t i f o r m nymphs. Developmental e f f e c t s o f kinoprene on s u r v i v i n g f i r s t i n s t a r a l a t i f o r m nymphs. S e q u e n t i a l l e t h a l e f f e c t s o f 65 ppm kinoprene a p p l i e d t o second i n s t a r a l a t i f o r m nymphs. D e v e l o p m e n t a l e f f e c t s o f 65 ppm k i n o p r e n e on s u r v i v i n g s e c o n d i n s t a r a l a t i f o r m nymphs. S e q u e n t i a l l e t h a l e f f e c t s o f 65 ppm kinoprene applied t o t h i r d i n s t a r a l a t i f o r m nymphs. ' Developmental e f f e c t s o f kinoprene on s u r v i v i n g t h i r d i n s t a r a l a t i f o r m .nymphs. ' '" > v  S e q u e n t i a l l e t h a l e f f e c t s o f 65 ppm k i n o p r e n e on f o u r t h i n s t a r a l a t i f o r m nymphs. Developmental e f f e c t s o f kinoprene on s u r v i v i n g f o u r t h i n s t a r a l a t i f o r m nymphs. O v e r a l l e f f e c t o f 10 ppm k i n o p r e n e on t h e f e c u n d i t y o f a p t e r i f o r m nymphs o f a p t e r o u s a d u l t s up t o 10 d a y s a f t e r a d u l t e m e r g e n c e . O v e r a l l e f f e c t o f 10 ppm k i n o p r e n e on t h e f e c u n d i t y o f a p t e r i f o r m nymphs o f a l a t e a d u l t s up t o 10 d a y s a f t e r a d u l t emergence.  vi  Page Table  7.  A p t e r a e p r o d u c t i o n by a p t e r o u s f e m a l e s w h i c h were t r e a t e d w i t h 65 ppm k i n o p r e n e w i t h i n 12 h r s . o f a d u l t emergence.  46  Table  8.  Effects of topical application o f p r e c o c e n e - I I on t h e f e c u n d i t y o f a p t e r i f o r m nymphs o f a p t e r o u s adults. E x c e p t on a p t e r o u s a d u l t s , k i n o p r e n e was a p p l i e d t o the precocene-treated i n s e c t s f o u r d a y s a f t e r a d u l t emergence.  48  Table  9.  Effects of topical application o f p r e c o c e n e - I I on t h e f e c u n d i t y o f a p t e r i f o r m nymphs o f a l a t e adults. K i n o p r e n e was a p p l i e d to the p r e c o c e n e - t r e a t e d i n s e c t s f o u r days a f t e r a d u l t emergence.  56  r  vii  L I S T OF FIGURES Page Figure  1.  C l i p - c a g e s used t o c o n f i n e C h i n e s e cabbage p l a n t .  a p h i d s on  12  D o r s a l view o f normal a l a t e a d u l t s showing t h e h e a v i l y s c l e r o t i z e d t h o r a x and d a r k l y p i g m e n t e d body.  20  D o r s a l view o f t h e head o f normal a l a t e a d u l t showing w e l l - d e v e l o p e d o c e l l i .  20  c.  Secondary antennal alate adult.  s e n s o r i a o f normal  20  d.  D o r s a l view o f normal a p t e r o u s a d u l t showing t h e absence o f s c l e r o t i z a t i o n i n the thorax.  20  D o r s a l view o f head o f normal a p t e r o u s a d u l t showing t h e absence o f o c e l l i .  20  T y p i c a l d e f o r m i t i e s produced i n malformed a l a t e a d u l t o i d s which had been t r e a t e d w i t h k i n o p r e n e a s f i r s t i n s t a r nymphs.  21  b.  D o r s a l view o f t h o r a x showing t h e e f f e c t on w i n g s a n d g e n i t a l i a .  21  c.  D o r s a l v i e w o f head c a p s u l e e f f e c t on o c e l l i .  21  d.  E f f e c t o f k i n o p r e n e on s e c o n d a r y a n t e n n a l sensoria.  21  D e f o r m i t i e s produced i n malformed a l a t e a d u l t o i d s w h i c h h a d been t r e a t e d w i t h k i n o p r e n e a s s e c o n d i n s t a r nymphs.  24  b.  M a g n i f i e d view o f wing rudiments,  24  c.  Dorsal effect  d.  E f f e c t o f k i n o p r e n e on s e c o n d a r y a n t e n n a l sensoria.  F i g u r e 2a.  F i g u r e 3a,  F i g u r e 4a.  view o f head c a p s u l e on o c e l l i .  showing t h e  showing  the  24 24  viii Page F i g u r e 5a.  D e f o r m i t i e s produced i n malformed a l a t e a d u l t o i d s w h i c h had been t r e a t e d w i t h k i n o p r e n e as t h i r d i n s t a r nymphs.  27  b.  M a g n i f i e d view of balloon-shaped wings.  27  c.  D o r s a l view o f head c a p s u l e showing t h e e f f e c t on o c e l l i .  27  d.  E f f e c t o f k i n o p r e n e on s e c o n d a r y antennal sensoria.  27  V e n t r a l view of malformed a l a t e a d u l t s w h i c h had been t r e a t e d w i t h k i n o p r e n e a s a f o u r t h i n s t a r nymph.  30  b.  D o r s a l view o f head c a p s u l e showing the e f f e c t on o c e l l i .  30  c.  E f f e c t o f k i n o p r e n e on s e c o n d a r y antennal sensoria.  30  F i g u r e 7.  F e c u n d i t y o f n o r m a l a p h i d s and t h o s e t r e a t e d w i t h 10 ppm k i n o p r e n e a s f i r s t i n s t a r a p t e r i f o r m larvae of apterous adults.  34  Figure  F e c u n d i t y o f normal a p h i d s and t h o s e t r e a t e d w i t h 10 ppm k i n o p r e n e a s second i n s t a r a p t e r i f o r m l a r v a e o f apterous adults.  35  F i g u r e 9.  F e c u n d i t y o f n o r m a l a p h i d s and t h o s e t r e a t e d w i t h 10 ppm k i n o p r e n e a s t h i r d i n s t a r a p t e r i f o r m larvae of apterous adults.  36  F i g u r e 10.  F e c u n d i t y o f n o r m a l a p h i d s and t h o s e t r e a t e d w i t h 10 ppm k i n o p r e n e a s f o u r t h i n s t a r apteriform larvae of apterous adults.  37  Figure  F e c u n d i t y o f n o r m a l a p h i d s and t h o s e 40 t r e a t e d w i t h 10 ppm k i n o p r e n e a s f i r s t i n s t a r a p t e r i f o r m larvae of a l a t e adults.  F i g u r e 6a.  8.  11.  F e c u n d i t y o f normal a p h i d s and t h o s e t r e a t e d w i t h 10 ppm k i n o p r e n e a s second i n s t a r a p t e r i f o r m l a r v a e o f alate adults. F e c u n d i t y o f normal a p h i d s and those t r e a t e d w i t h 10 ppm k i n o p r e n e a s t h i r d instar apteriform larvae of alate adults. F e c u n d i t y o f normal a p h i d s and t h o s e t r e a t e d w i t h 10 ppm k i n o p r e n e a s fourth i n s t a r apteriform larvae of alate adults. F e c u n d i t y o f normal a p h i d s and those t r e a t e d w i t h 65 ppm k i n o p r e n e a s newly-ecdysed apterous a d u l t s . F e c u n d i t y o f c o n t r o l a p h i d s and those ..with 0.5 ug p r e c o c e n e - I I a s f i r s t i n s t a r a p t e r i f o r m nymphs o f a p t e r o u s adults. The e f f e c t s o f 10 ppm kinoprene a p p l i e d t o precocenet r e a t e d i n s e c t s f o u r days a f t e r a d u l t emergence a r e a l s o i n d i c a t e d . F e c u n d i t y o f c o n t r o l a p h i d s and those t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I a s s e c o n d i n s t a r a p t e r i f o r m nymphs o f apterous adults. The e f f e c t s o f 10 ppm k i n o p r e n e a p p l i e d t o p r e c o c e n e t r e a t e d i n s e c t s f o u r days a f t e r a d u l t emergence a r e a l s o i n d i c a t e d . F e c u n d i t y o f c o n t r o l a p h i d s and t h o s e t r e a t e d w i t h 0.5 u g p r e c o c e n e - I I a s t h i r d i n s t a r a p t e r i f o r m nymphs o f apterous adults. The e f f e c t s o f 10 ppm k i n o p r e n e a p p l i e d t o p r e c o c e n e t r e a t e d i n s e c t s four days a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  F e c u n d i t y o f c o n t r o l a p h i d s and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I a s f o u r t h i n s t a r a p t e r i f o r m nymphs o f apterous adults. The e f f e c t s o f 10 ppm k i n o p r e n e a p p l i e d t o p r e c o c e n e t r e a t e d i n s e c t s f o u r days a f t e r a d u l t emergence a r e a l s o i n d i c a t e d . F e c u n d i t y o f c o n t r o l aphids and those t r e a t e d w i t h 0.2 ug p r e c o c e n e - I I a s newly-ecdysed apterous a d u l t s . The e f f e c t s o f 65 ppm k i n o p r e n e a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s 24 h r s . o r 4 days a f t e r a d u l t emergence a r e a l s o indicated. F e c u n d i t y o f c o n t r o l a p h i d s and those t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I a s f i r s t i n s t a r a p t e r i f o r m nymphs o f alate adults. The e f f e c t s o f 10 ppm kinoprene applied t o precocene-treated i n s e c t s f o u r days a f t e r a d u l t emergence are a l s o i n d i c a t e d . F e c u n d i t y o f c o n t r o l a p h i d s and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I a s s e c o n d i n s t a r a p t e r i f o r m nymphs o f alate adults. The e f f e c t s o f 10 ppm kinoprene applied t o precocene-treated i n s e c t s f o u r d a y s a f t e r a d u l t emergence are a l s o i n d i c a t e d . F e c u n d i t y o f c o n t r o l aphids and those t r e a t e d w i t h 0.5 u g p r e c o c e n e - I I a s t h i r d i n s t a r a p t e r i f o r m nymphs o f alate adults. The e f f e c t s o f 10 ppm kinoprene applied to precocene-treated i n s e c t s f o u r d a y s a f t e r a d u l t emergence are also indicated. F e c u n d i t y o f c o n t r o l aphids and those t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I a s f o u r t h i n s t a r a p t e r i f o r m nymphs o f alate adults. The e f f e c t s o f 10 ppm kinoprene applied t o precocene-treated i n s e c t s f o u r d a y s a f t e r a d u l t emergence are also i n d i c a t e d .  xi  ACKNOWLEDGEMENTS  I s i n c e r e l y t h a n k t h e f o l l o w i n g p e r s o n s who to the success  o f my  research:  - My s u p e r v i s o r , help,  D r . R.H. E l l i o t t  invaluable ideas,  stages  contributed  of this  for h i s unfailing  and encouragement i n a l l  work.  - Members o f my c o m m i t t e e ,  D r . R.H. E l l i o t t ,  D r . A.R.  F o r b e s , D r . J.H. M y e r s , a n d D r . V.C. R u n e c k l e s f o r their  suggestions  i n preparing  this  thesis.  - S p e c i a l t h a n k s t o C h o - k a i Chan a n d D r . A.R. F o r b e s o f t h e A g r i c u l t u r e Canada R e s e a r c h for  providing  a p h i d s and t a k i n g  Dr.  B.D. Frazer  provided  me w i t h  S t a t i o n , Vancouver,  the; e l e c t r o n m i c r o g r a p h s . clip-cages.  - To a l l my l a b . m a t e s , D a l e , E l n o r a , Susan who have b e e n v e r y - To my b r o t h e r - i n - l a w , their  Rosemary a n d  i n t e r e s t i n g and h e l p f u l f r i e n d s .  V i j a y , and s i s t e r ,  encouragement and c o - o p e r a t i o n  Shiksha, f o r  t h r o u g h o u t my  thesis. - L i s a Walker,  for her expert  t y p i n g and t i m e l y  help.  1  INTRODUCTION  The serious The  green  p e a c h a p h i d , Myzus p e r s i c a e  a g r i c u l t u r a l pest throughout  (Sulzer) i s a  the world  (Essig  s p e c i e s i s p o l y p h a g o u s and has been o b s e r v e d  host p l a n t s  (Forbes  p e r s o n a l cortimunication) .  1926).  on o v e r  200  The o v e r -  wintering  e g g s a r e l a i d . o n woody p e r e n n i a l - i n c l u d i n g p e a c h ,  nectarine  and o c c a s i o n a l l y a p r i c o t , s w e e t  Damage t o t h e w i n t e r infestations  cause  host  serious leaf  o f buds a n d / o r p r e m a t u r e the  formation  occurs  o f winged  fruit  lettuce,  and a v a r i e t y  Compared t o o t h e r p a t t e r n o f aphids conditions,  of  o r plum.  i n t h e s p r i n g when  heavy  damage, w i t h e r i n g a n d d r o p p i n g set.  forms which  summer h o s t p l a n t s i n c l u d i n g  cherry  These c o n d i t i o n s subsequently  crucifers,  favour  migrate t o  c u c u r b i t s , tomatoes,  ornamentals.  insect  species, the  i s exceedingly  complex.  t h e a d u l t f e m a l e may r e p r o d u c e  developmental Depending  upon  sexually -  -(oviparous) _6r - p a r t h e n o g e n e t i c a l l y ( v i v i p a r o u s )  \  j  .  (Blackman 1974) .  ( In a d d i t i o n , aphids species,  are polymorphic  several distinct  i n that, within a  morphological  forms a r e p r o d u c e d .  T h e s e f o r m s , c a l l e d morphs, have d i f f e r e n t cycle. fall,  Besides there  roles  t h e s e x u a l morphs, w h i c h o c c u r  are alate  and a p t e r o u s  v i r g i n o p a r a e do n o t d e v e l o p p r o d u c e a s many o f f s p r i n g 19 75). However, t h e a l a t e  single  i n the l i f e  only  virginoparae.  i n late  The a l a t e  as q u i c k l y a s t h e a p t e r o u s  forms o r  (Blackman 19 74; Mackay a n d W e l l i n g t o n condition provides  a means o f e s c a p e  2  from d e t e r i o r a t i n g food sources,. t h e r e b y e n a b l i n g the i n s e c t s e s t a b l i s h new as  c o l o n i e s on f r e s h h o s t p l a n t s .  In species  such  p e r s i c a e and t h e c a b b a g e a p h i d , B r e v i c o r y n e b r a s s i c a e *  ( W h i t e 19 68b), t h e a l a t e and a p t e r o u s m o r p h s c a n n o t d i s t i n g u i s h e d e x t e r n a l l y as f i r s t However, i n t h i r d  and s e c o n d i n s t a r  nymphs.  and f o u r t h i n s t a r s , p r e s u m p t i v e a l a t e s  be r e c o g n i s e d b y s w e l l i n g s on t h e meso- and Three t h e o r i e s have been advanced morphism i n a p h i d s . winged  be  The  first  can  meta-thorax.  t o e x p l a i n wing  theory postulates that  polythe  f o r m i s t h e n o r m a l c o n d i t i o n and t h e w i n g l e s s f o r m  develops secondarily. Klodnitsky Davidson  T h i s t h e o r y has b e e n e x p o u n d e d  (1912), T u r n e r and B a k e r  (1927),  Rivnay  (1937),  (1916), A c k e r m a n  and S h u l l  (1938).  by (1926),  These  a u t h o r s h a v e shown h i s t o l o g i c a l l y t h a t w i n g a n l a g e n s o r r u d i m e n t s a r e p r e s e n t i n a l l e m b r y o s and whether  first  i n s t a r nymphs  they are destined t o develop into alates or apterae.  These f i n d i n g s prompted  the authors t o suggest t h a t  e n v i r o n m e n t a l and/or p h y s i o l o g i c a l  certain  f a c t o r s favour the pro-  d u c t i o n o f t h e a p t e r o u s f o r m by i n h i b i t i n g t h e d e v e l o p m e n t wing rudiments.  T h i s h y p o t h e s i s was  J o h n s o n a n d B i r k . (1960) who that prior to b i r t h ,  of  further substantiated  demonstrated i n Aphis  by  craccivora  the wing anlagens of presumptive apterae  were l a r g e r t h a n t h o s e o f . p r e s u m p t i v e a l a t e s . An o p p o s i n g t h e o r y h a s b e e n p r o p o s e d w h i c h s u g g e s t s t h a t the  w i n g l e s s f o r m i s t h e n o r m a l c o n d i t i o n and t h e w i n g e d  develops secondarily.  T h i s t h e o r y was  p o s t u l a t e d by  form  White  * The t a x o n o m i c a u t h o r i t i e s f o r t h i s and o t h e r a p h i d s p e c i e s l i s t e d i n Appendix 6  to  3  (194 6) who  c o u l d n o t d e t e c t any w i n g a n l a g e n s i n h i s t o l o g i c a l  s e c t i o n s o f e m b r y o s and y o u n g nymphs o f t h e c h r y s a n t h e m u m aphid, Macrosiphoniella sanborni. t h a t t h e w i n g l e s s f o r m was  A l t h o u g h i t was  suggested  the normal c o n d i t i o n , White  was  u n a b l e t o d e t e r m i n e when f o r m a t i o n o f w i n g s b e g a n o r what factors influenced their  development.  According t o the t h i r d theory, the presence or o f w i n g r u d i m e n t s i n e m b r y o s may species.  T h i s was  absence  d e p e n d upon t h e s t r a i n o f t h e  d e m o n s t r a t e d by K i t z m i l l e r  (1950) who  found  w i n g r u d i m e n t s i n h i s t o l o g i c a l s e c t i o n s o f some c h r y s a n t h e m u m aphids but not i n o t h e r s . A number o f e n v i r o n m e n t a l f a c t o r s h a v e b e e n shown t o i n f l u e n c e wing development  i n aphids.  s p e c i e s and s t a g e o f d e v e l o p m e n t ,  Depending  upon t h e  crowding appears t o  wing f o r m a t i o n i n a t l e a s t t h r e e d i f f e r e n t ways. p e r f o r m e d on t h e p o t a t o a p h i d , M a c r o s i p h u m  influence  Experiments  euphorbiae  (Shull  192 8 ) ; c h r y s a n t h e m u m a p h i d , M a c r o s i p h o n i e l l a s a n b o r n i (Kitzmiller  1950); v e t c h a p h i d , Megoura v i c i a e  1 9 6 7 ) ; and p e a a p h i d , A c y r t h o s i p h o n p i s u m  (Lees  1961,  (Sutherland  1969)  have d e m o n s t r a t e d t h a t c r o w d i n g d u r i n g t h e a d u l t s t a g e f a v o u r s the production of alate o f f s p r i n g . under crowded  Apterous adults reared  c o n d i t i o n s , always produced a l a t e  offspring  whereas those a d u l t s r e a r e d i n i s o l a t i o n , produced ones.  apterous  I n c o n t r a s t , o t h e r s t u d i e s have s u g g e s t e d t h a t m a t e r n a l  crowding i s unimportant but that crowding during e a r l y s t a g e s h a s a more p r o n o u n c e d  nymphal  e f f e c t on a l a t e p r o d u c t i o n ( N e i l s  4  1912;  Shinji  1918;  1951; J o h n s o n Glen 1971). their  Wadley  1923;  and B i r k 1960;  R i v n a y 1937;  Noda 1 9 6 1 ;  Bonnemaison  Toba 1967;  A p h i d s r e a r e d under crowded  Dixon  conditions  and  during  f i r s t and s e c o n d i n s t a r n y m p h a l s t a g e s a l w a y s d e v e l o p e d  i n t o a l a t e s i r r e s p e c t i v e of conditions to which t h e i r were exposed.  mothers  O t h e r s t u d i e s h a v e shown t h a t c r o w d i n g h a s  i n f l u e n c e on w i n g development.  Insects born t o a l a t e  w h i c h were r e a r e d under e i t h e r crowded  or isolated  always developed i n t o apterae i r r e s p e c t i v e of  no  adults,  conditions,  conditions  p r e v a l e n t d u r i n g t h e i r n y m p h a l s t a g e s ( S u t h e r l a n d 1970;  Mackay  1977) . A c c o r d i n g t o White changes  (1965, 1968b), e n v i r o n m e n t a l l y i n d u c e d  i n aphid polymorphism  are mediated through the endocrine  system, p a r t i c u l a r l y the c o r p o r a a l l a t a s i t u a t e d b e h i n d t h e b r a i n and p r o d u c e  These g l a n d s a r e  j u v e n i l e hormone ( J H ) .  As t h e n a t u r a l J H h a s n o t b e e n i s o l a t e d o f JH i n m o r p h o g e n e s i s  (CA).  from a p h i d s , the  and r e p r o d u c t i o n c a n o n l y be  implied  by e s t a b l i s h i n g t h e e f f e c t s o f j u v e n i l e hormone a n a l o g u e s on d e v e l o p m e n t  ( W h i t e and Lamb 1 9 6 8 ) .  In these s t u d i e s ,  JHA h a v e b e e n a d m i n i s t e r e d e i t h e r b y d i r e c t t o p i c a l o r by r e a r i n g a p h i d s on J H A - t r e a t e d p l a n t s . b e e n p e r f o r m e d on a l a t i f o r m 1980), D o r a l i s fabae 1968a, Hrdy  1971;  1974;  Such  l a r v a e o f jyL_ v i c i a e  H a n g a r t n e r e t a l . 1 9 7 1 ) , M_;_ p e r s i c a e  M i t t l e r e t a l . 1 9 7 6 ) , and i^L e u p h o r b i a e  and P e r r o n 1 9 7 5 ) .  (JHA) the  application  s t u d i e s have  (Lees  (Von Dehn 1 9 6 3 ) , B^ b r a s s i c a e  role  1977,  (White (Tamaki  1973;  (Cloutier  The m o r p h o l o g i c a l r e s p o n s e s i n d u c e d by  JHA  5  i n these  s p e c i e s i n c l u d e d (1) t h e p r o d u c t i o n o f  nymphal i n s t a r s ,  (2) d e l a y e d m e t a m o r p h o s i s ,  (3) m a l f o r m e d  wings,  (4) a b n o r m a l c a u d a and  and/or  (6) an i n c r e a s e d p r o p o r t i o n o f a p t e r a e .  Mittler M.  e t a l . (197 6) o b s e r v e d  genetalia,  rarily.  Furthermore,  (5) r e d u c e d  but l a r v i p o s i t i o n  had  s t a n t i a t e d by s t u d i e s on  fewer  secondary  reduced  and d e v e l o p m e n t o f t h e o c e l l i antennal p l a c o i d organs  by JHA  antennal sensoria T h i s was  or suppressed  b u t had  ( L e e s 1977,  sub-  shown t h a t sclerotization  no g r o s s e f f e c t s on  1980).  However, m i c r o s -  a l a t e s than i n c o n t r o l s  As many o f t h e m o r p h o l o g i c a l f e a t u r e s  suppressed  are c h a r a c t e r i s t i c of a l a t e a d u l t s but  i m m a t u r e s and  apterous  a d u l t females  as b e i n g n e o t e n i c .  a d u l t s , many a u t h o r s r e g a r d  larval  This retention  j u v e n i l e c h a r a c t e r i s t i c s presumes t h e e x i s t e n c e o f h i g h t i t e r s d u r i n g embryonic and/or l a r v a l development.  of  JH  Additional  f o r t h i s h y p o t h e s i s comes f r o m s t u d i e s on a p t e r i f o r m  s t a g e s w h e r e JHA activity  not  apterous  That i s , they r e t a i n  c h a r a c t e r i s t i c s when r e p r o d u c t i v e l y m a t u r e .  support  the  o f t h e o r g a n s i n d i c a t e d t h a t t h e y were  l e s s numerous i n J H A - t r e a t e d  treatments  and  i n c r e a s e d tempo-  v i c i a e , w h e r e i t was  h i g h c o n c e n t r a t i o n s o f JHA  ( L e e s 1980).  of  sclerotization  and no o c e l l i when t h e y emerged as a d u l t s .  s m a l l e r and  addition,  the authors c l a i m t h a t a l a t i f o r m l a r v a e  treated w i t h kinoprene  copic examination  In  fecundity  t h a t when f o u r t h i n s t a r s  p e r s i c a e were t r e a t e d w i t h k i n o p r e n e ,  p i g m e n t a t i o n were reduced  supernumerary  h a v e b e e n shown t o e x h i b i t l i t t l e  (Hangartner  * secondary  e t a l . 1971;  morphogenic  C l o u t i e r and P e r r o n  antennal s e n s o r i a (antennal p l a c o i d  1975).  organs)  6  As d i s c u s s e d by M i t t l e r e t a l . ( 1 9 7 6 ) , t h e m o r p h o g e n i c effects  i n d u c e d by JHA  terms because l i t t l e  are d i f f i c u l t  to interpret  i s known r e g a r d i n g t h e n o r m a l  mediation of morphogenesis i n aphids. l e v e l o f JH i n t h e haemolymph h a s  Gilbert  1975).  endocrine  In o t h e r s p e c i e s , the  b e e n shown t o r e a c h i t s  l o w e s t l e v e l d u r i n g t h e f i n a l nymphal s t a g e 1970;  i n hormonal  (cf.  T h i s d e c l i n e i n JH t i t e r  Wigglesworth  enables  the  i n s e c t t o metamorphose i n t o t h e a d u l t s t a g e c h a r a c t e r i s e d by f u l l y developed  wings.  As  current techniques prevent  direct  measurement o f J H l e v e l s i n a p h i d s , i n d i r e c t a s s e s s m e n t s h a v e b e e n made on t h e b a s i s o f t h e h i s t o l o g i c a l CA was of  (White  1965;  1968b; E l l i o t t  1975).  In these  assumed t h a t c h a n g e s i n t h e v o l u m e and t h e CA  appearance o f  studies, i t  nuclear  diameters  r e f l e c t t h e p a t t e r n o f J H s y n t h e s i s and r e l e a s e .  T h a t i s , a l a r g e g l a n d w i t h e n l a r g e d n u c l e i was a small gland i n a c t i v e . Elliott  independently  believed active,  B a s e d on t h e s e a s s u m p t i o n s ,  suggested  t h a t t h e CA  White  i n apterous  i n alate  inactive.  i n a p t e r o u s morphs w o u l d  Thus, h i g h JH t i t e r s  l a r v a e were  t h e r e t e n t i o n o f j u v e n i l e c h a r a c t e r i s t i c s and However, i n a d u l t s , t h e CA  i n a c t i v e whereas those  active.  Conversely,  In the  in  latter  the apterous  a d u l t s , l o w JH l e v e l s w e r e p o s t u l a t e d t o h a v e l i t t l e the form of the o f f s p r i n g ,  favour  f o r m s became  c a s e , t h e h i g h e r JH l e v e l s w e r e b e l i e v e d t o f a v o u r production of apterous o f f s p r i n g .  relatively  stimulate ovarian  of apterous  i n a l a t e s became  and  larvae  w e r e v e r y a c t i v e w h e r e a s t h e CA  growth.  the  effect  so t h a t J H i n f l u e n c e s c o u l d  be  on  7  modified logical  p o s t n a t a l l y by  various  environmental  s t u d i e s appear t o c o n f i r m  this  hypothesis  (1968b) d e m o n s t r a t e d t h a t t h e mean n u c l e a r i n the  CA  of aphids  significantly  This  of the  Recently  (apteriforms)  insects reared  histological  t h a t the  criticism nuclear  for assessing  d i a m e t e r and glandular  u l t r a s t r u c t u r e of the  volume and  apterous-producing  a d u l t s were n o t  Furthermore, crowding, which CA  ultrastructure.  of Lees  CA  involved  i n the  viciae.  Rather, the  not  ment i n of  the  CA  alate  1976).  females  Since  from a l a t e -  contrary  to  i s tenable  maternal  develop-  only  amounts  adults  were i n a c t i v e i n  insect species  studied,  o v a r i a n development  Mackay and  had the  t h a t the  of apterous CA  and  ( L e c k s t e i n 1976).  presence of considerable CA  are  example,  a p t e r o u s f e m a l e s a r e more f e c u n d  (Blackman 1974,  CA  demonstrated  p r e n a t a l c o n t r o l o f wing  However, i n most o t h e r  Leckstein hypothesis  CA  (1967), L e c k s t e i n c o n c l u d e d  have b e e n shown t o s t i m u l a t e  Gilbert  have  alate production  prompted L e c k s t e i n t o p o s t u l a t e t h a t the insects!  Elliott  For  viciae  Therefore,  smooth e n d o p l a s m i c r e t i c u l u m i n t h e  these  hyper-  volume o f t h e  different  stimulates  suggestion was  were  authors  activity.  t h a t the  on  caused  i n that other  s t u d i e s p e r f o r m e d on  effect  cells  together  s t u d i e s o f W h i t e and  e l e c t r o n microscopy  no  White  CA.  the  poor c r i t e r i a  of  as  Histo-  diameters of  suggested t h a t i s o l a t i o n  have come u n d e r g r e a t established  in isolation  l a r g e r than those  (alatiforms). activity  reared  factors.  Wellington  (cf. than  1975),  the  i f some h o r m o n a l a g e n t .  8  o t h e r t h a n JH r e g u l a t e s r e p r o d u c t i v e d e v e l o p m e n t aphids.  As w i l l  be d i s c u s s e d s h o r t l y , t h i s  seems  i n female unlikely.  A second c r i t i c i s m o f t h e t h e o r y t h a t JH s u p p r e s s e s wing development  i n a p h i d s came f r o m s t u d i e s p e r f o r m e d on  Using a r t i f i c i a l t h a t t w o JHA  d i e t s , A p p l e b a u m e t a l . (1975) d e m o n s t r a t e d  ( c h l o r i n a t e d f a r n e s o i c a c i d and an i s o m e r i c  m i x t u r e ) had no a p p a r e n t p r e n a t a l o r p o s t n a t a l e f f e c t on development  persicae.  persicae.  apterizing  I n s t e a d , t h e compounds c a u s e d d e l a y e d  and h i g h m o r t a l i t y .  The  authors viewed these  e f f e c t s as p a t h o l o g i c a l and s u g g e s t e d t h a t e x o g e n e o u s  JHA  t r e a t m e n t s merely d i s r u p t the i n t e r n a l hormonal m i l i e u . m o r e , A p p l e b a u m e t a_l.  JH  Further-  suggested t h a t aphids are normally  p r e s u m p t i v e a p t e r a t e s w i t h dormant wing r u d i m e n t s w h i c h a r e l a t e r s t i m u l a t e d by p o s t n a t a l l i p i d As i n d i c a t e d a b o v e , is  i n doubt.  not  involved  fraction(s).  t h e r o l e o f JH i n a p h i d p o l y m o r p h i s m  No t h e o r y s t r o n g l y s u g g e s t s w h e t h e r i n wing development  and i f i t i s i n v o l v e d , when  and a t what s t a g e t h e hormone i s i m p o r t a n t . e v i d e n c e f o r and a g a i n s t hormonal in  aphids, Hales  JH i s o r i s  In reviewing the  c o n t r o l of wing  polymorphism  (1976) s u g g e s t e d t h a t t h e r o l e o f J H , i f a n y ,  c o u l d be d e m o n s t r a t e d o n l y w i t h new  techniques involving  u s e o f a n t i - j u v e n i l e hormone o r a n t i - a l l a t o t r o p i c  the  agents.  A n t i - a l l a t o t r o p i c a g e n t s w e r e d i s c o v e r e d b y B o w e r s and his  a s s o c i a t e s i n 197 6 when t h e y i s o l a t e d two  d e r i v a t i v e s from the bedding p l a n t , Ageratum In  preliminary  chromene haustonianum.  s t u d i e s , two d e r i v a t i v e s , p r e c o c e n e - I and  9  p r e c o c e n e - I I , were shown t o i n d u c e p r e c o c i o u s m e t a m o r p h o s i s and  p r e v e n t o v a r i a n development i n t h e milkweed bug, c o t t o n  stainer,  a p p l e maggot, and bean b e e t l e .  responses  t o the precocenes  application. inhibited  locust,  with precocene-II precocene-I  within  irreversible  fasciatus  (Pener  1978) l e d t o p r e c o c i o u s m e t a m o r p h o s i s .  (Schoonveld  o f precocene-I  i n the desert locust,  precocene-II  was t o t a l l y  comparable stages  In the  o f young nymphs  on t h e CA were  d e g e n e r a t i v e changes i n t h e g l a n d s  application  morphosis  (Unnithan e t a l . 1977).  e f f e c t s o f the precocenes  i n that  ( P r a t t and  et. a l . 1978; Nemec e t a l . 1978) o r  two h r s . o f t r e a t m e n t  topical  precocene-II  d e g e n e r a t i o n o f t h e CA  Locusta m i g r a t o r i a , treatment  (Pedersen  inhibitory  immediate  that  JH b i o s y n t h e s i s i n P e r i p l a n e t a a m e r i c a n a  of a d u l t Oncopeltus  The  v a r i e d w i t h d o s a g e and t i m e o f  F u r t h e r s t u d i e s demonstrated  Bowers 1 9 7 7 ) , and i n d u c e d  migratory  The e x t e n t o f t h e  1979).  caused  almost  began  Although  p r e c o c i o u s meta-  Schistocerca gregaria,  i n a c t i v e when a p p l i e d  ( P a q u i n and P e r r o n  topically to  1978; U n n i t h a n  e t a l . 1980).  However, when S ^ g r e g a r i a nymphs were e x p o s e d t o p r e c o c e n e - I I residues  i n petri-dishes  f o r 24 h r s . , a h i g h p r o p o r t i o n o f t h e  t r e a t e d nymphs metamorphosed p r e c o c i o u s l y ( U n n i t h a n T h u s , b i o a s s a y methods c a n s t r o n g l y action authors  o f the precocenes have s u g g e s t e d  when t h e CA a r e a c t i v e 1979).  e t a l . 1980).  influence the inhibitory  on CA a c t i v i t y .  t h a t t h e precocenes  Moreover, are only  (Masner e t a l . 1979; U n n i t h a n  several inhibitory and N a i r  10  To  d a t e , o n l y one  p r e c o c e n e on on  t h e pea  topical in  aphid  aphid,  s t u d y has  i n v e s t i g a t e d the  m o r p h o g e n e s i s and A^  of a l a t e o f f s p r i n g even though  formation  p r o d u c e d was  h i g h l y v a r i a b l e and  of apterae.  was  applied.  has  an a n t i a l l a t o t r o p i c  The  as  i n M_j_  reproduction,  in  uncertainty  persicae,  structures.  e f f e c t i n aphids,  f e m a l e s was  the  reversed  in  somethe  aphids. of the  k i n o p r e n e and  (2) w i n g d e v e l o p m e n t , and  precocene-II  However, as t h e d e c l i n e  chemical probes t o e s t a b l i s h the  metamorphosis,  alates  a p p l i c a t i o n s , i t would appear t h a t  stimulate reproduction  genesis  number o f  (alate-inducing)  l a r v i p o s i t i o n i n precocene-treated  In l i g h t of the  resulted  d e p e n d e d upon when p r e c o c e n e  cause s t e r i l i t y .  s u b s e q u e n t JH  that  conditions  Although such e f f e c t s suggested t h a t  compound d i d n o t  may  Working  p i s u m , M a c k a u e r e t a l . (1979) showed  favoured the  CA  reproduction.  a p p l i c a t i o n of precocene-II to apterous adults  the production  w h a t by  e f f e c t s of  r o l e o f JH  i n morpho-  p r e c o c e n e - I I were used  i n f l u e n c e o f JH o n (3)  (1)  sclerotization,  (5) d i f f e r e n t i a t i o n o f c e p h a l i c  sensory  (4)  11  MATERIAL AND METHODS  A.  REARING  TECHNIQUES  Green peach a p h i d s were o b t a i n e d  from Dr. Forbes  a t t h e A g r i c u l t u r e Canada R e s e a r c h S t a t i o n , V a n c o u v e r . a p h i d s were r e a r e d The  on C h i n e s e c a b b a g e , B r a s s i c a e  p l a n t s w e r e t r a n s f e r r e d i n t o mesh c a g e s  The  chinensis_.  ( 3 6 x 4 0 x 3 9 cm.)  l o c a t e d i n t h e Department o f P l a n t S c i e n c e greenhouse. stock  The  c o l o n i e s w e r e m a i n t a i n e d a t 1 6 ° - 18° C a n d 16L:8D  photoperiod.  To e n s u r e v i g o r o u s  were r e p l a c e d  as they  (a)  c o l o n i e s , cabbage  plants  deteriorated.  A l a t i f o r m nymphs For  s t u d i e s r e q u i r i n g a l a t i f o r m nymphs, n e w l y -  e c d y s e d nymphs w e r e c o l l e c t e d f r o m s t o c k virginoparae  which had been r e a r e d  colonies o f apterous  under crowded  conditions.  To. e n s u r e c o n t i n u o u s c r o w d i n g , .45 i n s e c t s w e r e t r a n s f e r r e d into a clip-cage  (3.2 cm. d i a m e t e r ) w h i c h was a t t a c h e d  l e a f o f a 3-week-old cabbage p l a n t placed  to a  ( F i g . 1 ) . The p l a n t s  were  i n a P e r c i v a l g r o w t h chamber a n d m a i n t a i n e d a t c o n d i t i o n s  described  above.  When t h e a p p r o p r i a t e  nymphal o r a d u l t  s t a g e s had been r e a c h e d , t h e i n s e c t s were t r e a t e d w i t h t h e t e s t compound d e s c r i b e d (b)  B(a).  Apteriform  nymphs a n d a d u l t s  Apteriform  nymphs w e r e o b t a i n e d  alate virginoparae order  i n Section  t o maintain  w h i c h had been r e a r e d isolation,  from apterous and i nisolation.  t h e nymphs w e r e r e a r e d  In  singly  12  13  i n c l i p - c a g e s (Mackauer e t a l . 1979).  The  a p p r o p r i a t e nymphal  o r a d u l t s t a g e s w e r e t r e a t e d w i t h t e s t compound d e s c r i b e d i n Section B . (c)  Alate-producing apterous  r e a r i n g newly-ecdysed apterous (20  insects/cage).  Kinoprene  a d u l t s were o b t a i n e d  a d u l t s i n crowded c o n d i t i o n s was  a p p l i e d to these  w i t h i n 12 h r s . o f t h e i r e m e r g e n c e .  To m a i n t a i n  The  B.  caged together  o f f s p r i n g were examined  Kinoprene The  (40 i n s e c t s / c a g e ) .  daily.  treatments  j u v e n i l e hormone a n a l o g u e ,  kinoprene  (2E,4E)-3,7,ll-trimethyl-2,4-dodecadienoate; Chem. C o r p . ) , w h i c h has i n VL_ p e r s i c a e  (2-propynyl  ZR-777;  Zoecon  b e e n shown t o be b i o l o g i c a l l y  ( M i t t l e r e t a l . 1976;  i n d i s t i l l e d w a t e r c o n t a i n i n g 0.1%  B o y d 1979), was  active prepared  of the w e t t i n g agent,  Tween  (polyoxyethylene s o r b i t a n monolaurate). The  l e t h a l and m o r p h o l o g i c a l e f f e c t s o f  w e r e e x a m i n e d on a l l n y m p h a l and and  produced  EXPERIMENTAL COMPOUNDS (a)  20  insects  crowded  c o n d i t i o n s among t h e i r p r o g e n y , a l l t h e o f f s p r i n g d a i l y w e r e c o l l e c t e d and  by  a l a t i f o r m morphs.  s t a g e s were d i p p e d  kinoprene  a d u l t stages of a p t e r i f o r m  W i t h i n 2 h r s . of m o l t i n g , the  i n a 0.1%  a q u e o u s Tween 20  t a i n i n g varying concentrations of kinoprene. t r e a t e d i n a s i m i l a r manner e x c e p t  various  solution  con-  C o n t r o l s were  t h e y were d i p p e d  in a  0.1%  14  a q u e o u s Tween 20 various  solution.  D e p e n d i n g upon t h e  t r e a t m e n t s were r e p l i c a t e d t h r e e  study,  the  to s i x times.  Studies  on  (1) a l a t i f o r m morphs i n c l u d e d 45 a p h i d s / r e p l i c a t e , (2)  on  i s o l a t e d a p t e r i f o r m nymphs and  r e p l i c a t e and  (3) t h o s e on  a d u l t s i n c l u d e d one  crowded a p t e r o u s a d u l t s  those  aphid/  included  20 i n s e c t s / r e p l i c a t e . The  e f f e c t s o f k i n o p r e n e on m o r t a l i t y , m e t a m o r p h o s i s ,  s c l e r o t i z a t i o n , wing formation, of the o c e l l i daily  f o r up (b)  and  secondary antennal  t o 10  days f o l l o w i n g  Precocene-kinoprene The  f e c u n d i t y and  differentiation  s e n s o r i a were  assessed  treatment,  treatments  a n t i - j u v e n i l e hormone compound,  precocene-II  ( 6 , 7 - d i m e t h y l - 2 , 2 - d i m e t h y l - c h r o m e n e ; A l d r i c h Chem. Co.) s t o r e d u n d e r n i t r o g e n a t 6°  C.  In p r e l i m i n a r y t e s t s ,  were exposed t o p r e c o c e n e v i a the e t a l . 1976).  This technique  fumigation  method  i n v o l v e s c o a t i n g the  was aphids  (Bowers bottom  of a glass p e t r i - d i s h w i t h a precocene-acetone s o l u t i o n , a l l o w i n g the acetone to evaporate then i n t r o d u c i n g the i n t o the p e t r i - d i s h f o r v a r y i n g bioassay  has  intervals.  b e e n u s e d s u c c e s s f u l l y on o t h e r  Although species  e t a l . 1980) , t h e m e t h o d p r o v e d i n a p p r o p r i a t e on because of i n c o n s i s t e n t e f f e c t s . r e f l e c t one it  This  inconsistency  major shortcoming of fumigation  i s impossible  to administer  precocene to each i n s e c t .  To  a uniform accomplish  aphids  the (Unnithan persicae may  i n that  d e f i n a b l e dose  of  this,  was  a method  15  developed  for topically  precocene-II al.  1979).  s o l u t i o n s were p r e p a r e d However,  t h i s proved  a c e t o n e was v e r y t o x i c problem  was  sulfoxide  a p p l y i n g precocene.  Initially,  i n acetone  unsatisfactory  (Mackauer e t because  the  t o the aphids v i z - 9 0 % m o r t a l i t y .  a v o i d e d by d i s s o l v i n g p r e c o c e n e  This  i n dimethyl  (DMSO). S o l u t i o n s o f precocene-DMSO were drawn i n t o  a 100 u l  H a m i l t o n m i c r o - s y r i n g e w h i c h was mounted on a m i c r o - a p p l i c a t o r ( I n s t r u m e n t a t i o n S p e c i a l t i e s Comp.). tration  o f precocene  method e n a b l e d me precocene adult  0.5  ug p r e c o c e n e  However, when t h i s  few by  totally  offspring  were t r e a t e d was  d o s e was  produced.  aphid.  I n most nymphal a n d  was a p p l i e d  t o each  insect.  administered to apterous  This d i f f i c u l t y  ug p r e c o c e n e  was  adults, i t  toxic  t o the  circumvented  to apterous adults.  i n an i d e n t i c a l manner e x c e p t  Controls  t h a t o n l y DMSO  applied. After  clip-cages chamber. daily  treatment,  t h e i n s e c t s were t r a n s f e r r e d  a n d p l a c e d on c a b b a g e p l a n t s The l e t h a l  65 ppm  i n a Persival  a n d m o r p h o l o g i c a l e f f e c t s were  f o r up t o 10 d a y s .  emergence, h a l f with  concen-  t h e volume a p p l i e d , t h e  prevented reproduction or proved  a p p l y i n g 0.2  the  t o a p p l y m i n u t e b u t p r e c i s e amounts o f  t o t h e abdomen o f e a c h  tests,  nearly  or adjusting  By a l t e r i n g  In a d d i t i o n ,  4 days a f t e r  growth  observed adult  o f t h e p r e c o c e n e - t r e a t e d i n s e c t s were  kinoprene.  into  treated  16  (c)  Statistical  analysis  Statistical  differences  kinoprene-precocene-treated two sample mean t e s t of  Where:  g r o u p s were d e t e r m i n e d u s i n g t h e  a t t h e 5% s i g n i f i c a n c e l e v e l . T h e  T was d e t e r m i n e d u s i n g  the following  X,-  t  *2  insects  : Xj= mean o f t r e a t e d  insects  of r e p l i c a t e s i n control  insects  n= number o f r e p l i c a t e s i n t r e a t e d  insects  m  > e r  Sp= p o o l e d  and  studies control  testedusing Programe).  value  formula  X,= mean o f c o n t r o l  , n= n u t  In  between t h e c o n t r o l and  variance  i n v o l v i n g .precocene - t r e a t e d , i n s e c t s , mean d i f f e r e n c e Duncans  Multiple  Range  kinoprene-treated  a t t h e 5% l e v e l  were  t e s t ( U . B . C . M F A V Computer  17  RESULTS  A.  KINOPRENE (a)  TREATMENTS  First  instar  When f i r s t in  a solution  than  apparent  alatiform  directly  adults,  a l a t i f o r m nymphs i n t h e k i n o p r e n e supernumerary molt observed By  (Table l b ) .  a t the molt  t h e time  w i t h kinoprene had d i e d . instar  nymphs d e v e l o p e d  into apterae. in  (representing  three r e p l i c a t e s .  Although,  the apterous i n s e c t s Two d a y s a f t e r was o b s e r v e d produced  treatments  controls,  underwent a  Significant mortality  i n t o normal  instar  stadium. treated  9.6%  a l a t e s were  developed produced  I n s t e a d , t h r e e malformed p o p u l a t i o n ) and  (2.2%) were p r o d u c e d  m o l t i n g was s l i g h t l y  a d u l t emergence, v i r t u a l l y aphids.  i n the  delayed,  d i d n o t undergo a supernumerary  i n the treated  was  85.1% o f t h e f i r s t  alates,  no n o r m a l  apterae  sub-  87.3% o f t h e s u r v i v i n g  2.2% o f t h e o r i g i n a l  t h r e e a p p a r e n t l y normal  instar  In t h e c o n t r o l s ,  In c o n t r a s t ,  greater  and e a c h  95.4% o f t h e nymphs  the k i n o p r e n e - t r e a t e d group.  alates  instar  and d u r i n g t h e f i f t h  of adult ecdysis,  effects  The % m o r t a l i t y  Unlike the fourth into  lethal  dipped  was s i g n i f i c a n t l y  during the f i r s t  nymphal s t a g e .  which molted  nymphs were  (Table l a ) .  i n t h e J H A - t r e a t e d group  i n the controls  sequent  instar  nymphs  c o n t a i n i n g 65 ppm k i n o p r e n e ,  were i m m e d i a t e l y observed  alatiform  molt.  100% m o r t a l i t y  No o f f s p r i n g  were  by t h e k i n o p r e n e - t r e a t e d a p h i d s d u r i n g t h i s  period  18  Table l a .  Sequential applied  lethal effects  to f i r s t  instar  o f 65 ppm  alatiform  Numbers i n p a r e n t h e s e s r e p r e s e n t mortality  values.  nymphs. cumulative  I n t h i s and r e m a i n i n g  means on t h e same row f o l l o w e d by superscripts at 45  are not s i g n i f i c a n t l y  t h e 5% l e v e l  tables,  similar different  (3 r e p l i c a t e s / t r e a t m e n t ,  insects/replicate). %  Stage  1st  kinoprene  mortality  Control  instar  2.2±1.3  Kinoprene-treated  3.7±1.5  a  b  2nd i n s t a r  1.5±0.8 (3.7±1.5)  3.0±2.0  b  (6.6±1.3)  3rd  instar  1 . 5 ± 0 . 8 ( 3 . 2 + 1.9)  3.0±0.7  b  (9.6±2.0)  4th  instar  0 . 0 ± 0 . 0 ( 5 . 2 + 1.9)  a  a  a  0.7±0.8 (10.3+2.0) b  Supernumerary m o l t  N.A.  41.614.1 !(51.9±3.9)  Supernumerary  N.A.  43.6±3.9  instar  Adult  (2-day-old)  Table  lb.  0.0±0.0 (5.2±1.9) a  Developmental e f f e c t s first  instar  Supernumerary  instars  Malformed a l a t e s Normal  alates  Normal  apterae  b  surviving  nymphs.  % original Stage  4.4+0.0 (99.8±0.0)  o f k i n o p r e n e on  alatiform  (95.4±0.0)  population  Control  Kinoprene-treated  0±0  87.3±0.7  0±0  2.2±1.3  85.113.9 9.612.0  0.010.0 a  2.2±1.3  b  19  whereas the  controls  produced  Before describing kinoprene, some o f  i t would  the  2-4  the  offspring/day.  aberrations  seem a p p r o p r i a t e  important morphological  n o r m a l a l a t e and  apterous adults.  induced  to b r i e f l y features  As  describe  exhibited  illustrated  scanning e l e c t r o n photomicrographs ( F i g s . 2a-c), typical  of  alate adults  thorax,  darkly  include  are  p i g m e n t e d body, w e l l - d e v e l o p e d  characterized  p i g m e n t e d body 2e).  ( F i g . 2d),  I n b o t h morphs, t h e  by  reduced  and  in  by  the  features  a heavily sclerotized  numerous s e c o n d a r y a n t e n n a l s e n s o r i a . adults  by  the  In  and  contrast,  apterous  sclerotization,  absence of  c a u d a and  ocelli  and  ocelli  genital plate  an  un-  (Fig.  are  prominent. The  kinoprene-treated  abnormal a d u l t o i d s , and  features  i n s e c t s , which molted  e x h i b i t e d many j u v e n i l e c h a r a c t e r i s t i c s  intermediate  between a l a t e and  Compared t o n o r m a l a l a t e s , t h e y had (Fig. fourth  i n s t a r a l a t i f o r m nymphs.  unpigmented.  the  Although o c e l l i  a p p e a r e d a b n o r m a l and The  t h o r a x was  3d)  whereas the  Like  wings  fourth  instars  unsclerotized  and  (Fig. 3c),  were s h r u n k e n and  p r e s u m p t i v e c a u d a and in  appearance.  in  and  the  l e s s developed than i n a l a t e  were more nymphal t h a n a d u l t  adults.  rudiments found  were p r e s e n t  secondary antennal s e n s o r i a  (Fig.  apterous  g r e a t l y reduced  3a, b) w h i c h .more c l o s e l y r e s e m b l e d  apterous adults,  into  cuticle they  adults. deformed  genital  plate  20  Fig.  2a.  Dorsal  view o f normal a l a t e  heavily body b.  adult  s c l e r o t i z e d t h o r a x and d a r k l y  Dorsal  pigmented  view o f t h e head o f normal a l a t e  Secondary antennal adult  d.  the  (2 3 X ) .  showing w e l l - d e v e l o p e d o c e l l i c.  showing  sensoria  adult  (187 X ) .  o f normal  alate  (937 X ) . •  Dorsal  view o f normal a p t e r o u s a d u l t  showing  the  absence o f s c l e r o t i z a t i o n i n the t h o r a x  (65  X).  Dorsal showing  view o f head o f normal a p t e r o u s t h e absence o f o c e l l i  (187 X ) .  adult  21  Fig.  3a.  Typical  deformities present  i n malformed  a d u l t o i d s w h i c h had been t r e a t e d w i t h as b.  first  and g e n i t a l i a  D o r s a l view o f head on o c e l l i  d.  nymphs  D o r s a l view o f thorax wings  c.  instar  Effect  kinoprene  (23 X ) . showing t h e e f f e c t  on  (187 X ) . capsule  showing t h e  effect  (187 X ) .  of kinoprene  sensoria  alate  (1873 X ) .  on s e c o n d a r y  antennal  22  (b)  Second i n s t a r a l a t i f o r m  nymphs  When k i n o p r e n e was a p p l i e d f o r m nymphs, t h e i n i t i a l the p r e v i o u s study instar, and  (Table 2a).  insects.  instar  alati-  m o r t a l i t y was n o t a s h i g h a s i n By t h e e n d o f t h e f o u r t h  8-9% m o r t a l i t y was o b s e r v e d  control  t o second  i n the kinoprene-treated  Before molting i n t o adults,  81.4% o f  t h e J H A - t r e a t e d a p h i d s underwent a supernumerary molt 2b).  About  10% m o r t a l i t y  t h a t by t h e t i m e  of adult  i n s e c t s had d i e d .  into apterae.  and  emergence, 18.4% o f t h e t r e a t e d  i n t o normal  80.7% o f t h e s e c o n d alates,  11% d e v e l o p e d  As i n t h e p r e v i o u s experiment,  a l a t e s were p r o d u c e d stead,  o c c u r r e d d u r i n g t h i s s t a d i u m so  In the controls,  i n s t a r nymphs d e v e l o p e d  no  normal  i n t h e k i n o p r e n e - t r e a t e d group.  71.8% o f t h e i n s e c t s d e v e l o p e d  9.6% i n t o n o r m a l  apterae.  A l t h o u g h m o l t i n g was  t h e k i n o p r e n e - t r e a t e d nymphs w h i c h m o l t e d  apterous  adults  and  alates slightly  into  d i d n o t undergo a supernumerary molt.  (65%) was o b s e r v e d w i t h i n  by t h e f o u r t h  2 days o f a d u l t  day, a l l i n s e c t s h a d d i e d  group.  During t h i s period,  treated  a p h i d s whereas c o n t r o l s Kinoprene-treated  malformed a d u l t o i d s , Again,  In-  i n t o malformed  delayed,  mortality  (Table  emergence  i n the treated  no o f f s p r i n g were p r o d u c e d reproduced  nymphs a n d a p t e r o u s  t h e w i n g s were u n d e v e l o p e d  by  normally.  i n s e c t s , which molted  resembled  High  and t h e c u t i c l e  into adults. remained  23  Table  2a.  Sequential applied  lethal effects  to second i n s t a r  o f 65 ppm k i n o p r e n e alatiform %  Stage  nymphs.  mortality  Control  Kinoprene-treated  2nd  instar  2.9±1.9  4.4±1.3  3rd  instar  2.2±0.0 (8.0±1.6)  2.9±0.8 (  7.3±0.8)  4th  instar  2.9±0.7 (8.0±1.6)  1.5±0.8 (  8.811.3)  Supernumerary molt  N.A.  O.OiO.O  ( 8.8±1.3)  Supernumerary  N.A.  (8.Oil.6)  9.712.0  (18.5±1.5)  a  a  a  instar  b  a  b  Adult  (2-day-old)  O.OiO.O  (8.Oil.6)  65.8±8.1  (84.316.6)  Adult  (4-day-old)  O.OiO.O  (8.0±1.6)  15.516.6  (99.810.0)  Table  2b.  Developmental e f f e c t s second i n s t a r  o f k i n o p r e n e on  alatiform  nymphs.  % original Stage  Supernumerary  Control  instars  Malformed a l a t e s  surviving  population Kinoprene-treated  0+0  81.4+2.0  010  71.812.0  Normal  alates  80.712.0  Normal  apterae  11.011.3  0.0+0.0 a  9.610.8  b  24  Fig.  4a.  D e f o r m i t i e s produced i n malformed a d u l t o i d s which had been t r e a t e d  alate with  k i n o p r e n e as s e c o n d i n s t a r nymphs (23 X ) . b.  M a g n i f i e d view of wing rudiments  c.  D o r s a l v i e w o f head c a p s u l e showing e f f e c t on o c e l l i  d.  (93 X ) . the  (187 X ) .  E f f e c t o f k i n o p r e n e on s e c o n d a r y a n t e n n a l sensoria  (2810 X ) .  25  unsclerotized ocelli  and unpigmented  were c o m p l e t e l y a b s e n t  sensoria  deformed  developed f u l l y , s t u d y on f i r s t (c)  ( F i g . 4a,b).  In addition,  ( F i g . 4c) a n d t h e a n t e n n a l  ( F i g . 4 d ) . A l t h o u g h t h e cauda had n o t i t was more p r o m i n e n t t h a n i n t h e p r e v i o u s  instars.  Third  i n s t a r a l a t i f o r m nymphs  The l e t h a l a n d m o r p h o l o g i c a l e f f e c t s o f k i n o p r e n e on t h i r d the  i n s t a r nymphs a r e shown i n T a b l e s 3a a n d 3b.  c o n t r o l s w h i c h m o l t e d n o r m a l l y and d e v e l o p e d i n t o  primarily alate adults,  86.7% o f t h e t r e a t e d  went a s u p e r n u m e r a r y m o l t .  controls, mortality rose abruptly  just after adult day.  insects  However, c o m p a r e d t o i n the treated  emergence and a l l i n s e c t s d i e d  During t h i s period  under-  U n t i l t h i s t i m e , k i n o p r e n e had  no s i g n i f i c a n t e f f e c t on m o r t a l i t y . the  Unlike  insects  by t h e s i x t h  no o f f s p r i n g w e r e p r o d u c e d . A l -  t h o u g h no n o r m a l a l a t e s w e r e p r o d u c e d i n t h e J H A - t r e a t e d g r o u p , 6.7% o f t h e i n s e c t s  i n the three r e p l i c a t e s developed  into apterae a f t e r undergoing a s l i g h t delay i n molting. the  In  c o n t r o l s , a b o u t 80% o f t h e t h i r d i n s t a r nymphs d e v e l o p e d  i n t o normal a l a t e s  and 11% d e v e l o p e d i n t o a p t e r a e .  Kinoprene-treated i n s e c t s , which molted i n t o malformed a d u l t o i d s ,  r e s e m b l e d nymphs a n d a p t e r o u s a d u l t s .  c u t i c l e remained u n s c l e r o t i z e d The g e n i t a l p l a t e  and unpigmented  (Fig.  and cauda were a l s o w e l l - f o r m e d .  The  5a).  Although  26  Table  3a.  Sequential applied  l e t h a l e f f e c t s o f 65 ppm k i n o p r e n e  to third instar alatiform  % Stage  nymphs.  mortality  Control  Kinoprene-treated  3rd  instar  5.9±1.9  4th  instar  3.0±0.7 (8.9±1.3)  1.5i0.7 (  N.A.  O.OiO.O  ( 6.6il.3)  5.1il.3  a  a  Supernumerary  molt  a  6.6il.3)  b  Adult  (3-day-old)  O.OiO.O  (8.9±1.3)  73.2i3.4  (79.8±2.5)  Adult  (6-day-old)  O.OiO.O (8.9+1.3)  19.9±2.5  (99.7i0.7)  Table  3b.  D e v e l o p m e n t a l e f f e c t s o f k i n o p r e n e on third  instar alatiform  surviving  nymphs.  % o r i g i n a l population Stage  Supernumerary  Control  instars  Malformed a l a t e s  Kinoprene-treated  OiO  86.7i2.2  N.A.  86.7i2.2  Normal a l a t e s  80.0i2.2  Normal a p t e r a e  11.0±1.3  O.OiO.O a  6.7il.3  b  27  Fig.  5a.  D e f o r m i t i e s produced i n malformed a d u l t o i d s w h i c h had b e e n t r e a t e d k i n o p r e n e as t h i r d  b.  alate with  i n s t a r nymphs ( 2 3 X ) .  M a g n i f i e d view of balloon-shaped wings  (74 X ) . c.  D o r s a l v i e w o f head c a p s u l e showing e f f e c t on o c e l l i  d.  the  (187 X ) .  E f f e c t o f k i n o p r e n e on s e c o n d a r y a n t e n n a l sensoria  (2810 X ) .  9LT-A  28  w i n g s were s l i g h t l y  r e d u c e d compared t o n o r m a l a l a t e s ,  were b a l l o o n - s h a p e d a n d f l u i d - f i l l e d v e n a t i o n was a p p a r e n t . not  ( F i g . 5 b ) . Some w i n g  O c e l l i were p r e s e n t  as d e v e l o p e d as i n normal a l a t e s .  antennal  sensoria  (d)  were d e f o r m e d  (Fig. 5d).  to fourth  i n s t a r nymphs,  l e s s p r o n o u n c e d e f f e c t s on m o r t a l i t y ,  and  reproduction  and  4b).  than i n e a r l i e r  controls  (Table  molted d i r e c t l y was d e l a y e d  4a), nearly  into adults  slightly  fourth  into apterae.  into apparently  days a f t e r a d u l t lethal  effect.  during  4b).  molted  than  insects ecdysis  6-7 h r s . s o o n e r  In the controls,  94.5% o f  i n t o n o r m a l a l a t e s , 0.7%  In the t r e a t e d normal a l a t e  insects,  adults,  78.5% d e -  a b o u t 12% i n t o Up t o two  emergence, k i n o p r e n e h a d no s i g n i f i c a n t After  this,  mortality  rose  period.  s i g n i f i c a n t l y as  and a l l i n s e c t s had d i e d  No s i g n i f i c a n t e f f e c t on r e p r o d u c t i o n this  the fourth  greater  However,  a n d 3.7% i n t o n o r m a l a p t e r a e .  compared t o c o n t r o l s day.  ( T a b l e 4a  observed during  as c o n t r o l a d u l t s  i n s t a r nymphs d e v e l o p e d  malformed a l a t e s ,  instars  90% o f t h e t r e a t e d  (Table  insects.  kinoprene  morphogenesis,  i n s e c t s was s i g n i f i c a n t l y  than k i n o p r e n e - t r e a t e d  developed  treated  Although the % m o r t a l i t y  i n s t a r o f the treated  veloped  but  In addition, the  had  the  (Fig. 5c),  F o u r t h i n s t a r a l a t i f o r m nymphs When a p p l i e d  the  they  by t h e s e v e n t h  was o b s e r v e d  29  Table  4a.  Sequential lethal on  fourth instar  e f f e c t s o f 65 ppm alatiform  kinoprene  nymphs.  % mortality  4th  Stage  Control  instar'  4.4±1.3  Kinoprene-treated  5.9±2.0  a  b  Adult  (3-day-old)  0.010.0  1.5±0.7(7.4±1.4)  Adult  (4-day-old)  O.OiO.O  9 2 . 7 ± 1 . 4 (100±0.0)  Table  4B.  Developmental e f f e c t s fourth instar  o f kinoprene  alatiform  Malformed  alates  Normal  alates  Normal  apterae  surviving  nymphs.  % original Stage  on  Control  population Kinoprene-treated  0±0  11.8+4.1  94.8±0.7  a  78.5±4.9  b  0.7±0.7  a  3.7±0.7  b  30  Fig.  6a.  Ventral  view o f malformed  alate  adults  w h i c h had b e e n t r e a t e d w i t h k i n o p r e n e as a b.  c.  fourth  i n s t a r nymph  (32 X ) .  D o r s a l v i e w o f head c a p s u l e  showing  the  effect  on o c e l l i  Effect  o f k i n o p r e n e on s e c o n d a r y a n t e n n a l  sensoria  (187 X ) .  (1873 X ) .  31  U n l i k e , e a r l i e r nymphal s t a g e s , l e s s p r o n o u n c e d e f f e c t s on Although a l l the  unpigmented  t r e a t e d had  normal s i z e d wings w i t h  ( F i g . 6a).  still  remained  and  w i n g s became  o c e l l i were w e l l d e v e l o p e d  normal a l a t e s , t h e i r s e n s o r i a  fully  unsclerotized  A f t e r f i v e days, the  p o r t i o n o f t h e w i n g s d r i e d up Although the  had  f o u r t h i n s t a r a l a t i f o r m nymphs.  developed cauda, t h e i r c u t i c l e and  kinoprene  terminal wrinkled.  ( F i g . 6b)  as  were s l i g h t l y deformed  in (Fig.  6c) . (e)  Apteriform  nymphs o f a p t e r o u s  adults  P a r a l l e l e x p e r i m e n t s w e r e a l s o c o n d u c t e d on a l l four As  i n s t a r s of apteriform  larvae born to apterous  i n the p r e v i o u s s t u d i e s , the  d i p p e d i n 65 ppm  kinoprene.  o c c u r r e d w i t h i n 24 effects,  hrs.  i n s e c t s were  To  9 0%  avoid  mortality these  subsequent e x p e r i m e n t s were p e r f o r m e d u s i n g  kinoprene.  L i k e the  controls, a l l kinoprene-treated  developed i n t o normal apterous a d u l t s . the  initially  However, n e a r l y  after dipping.  adults.  controls, ecdysis  was  s e c o n d i n s t a r nymphs and  t h i r d and  fourth  i n s t a r nymphs.  a l a t i f o r m nymphs, a p t e r i f o r m  about 8 h r s . Unlike  10  ppm  insects  However, c o m p a r e d  d e l a y e d a b o u t 1 day  f i r s t and  lethal  in in  to  treated treated  kinoprene-treated  nymphs d i d n o t  undergo  any  t r e a t m e n t s on  fecundity  of  supernumerary m o l t s . The  e f f e c t s of the  the  32  e m e r g e n t f e m a l e s were a l s o a s s e s s e d .  O v e r a l l , no  significant  d i f f e r e n c e s were o b s e r v e d b e t w e e n t h e c o n t r o l s and k i n o p r e n e treated  insects  (Table  p r o d u c e d an a v e r a g e ment p e r i o d .  5).  Each  female i n b o t h  3.2 o f f s p r i n g / d a y  In t o t a l ,  fecundity all  The d a i l y o f the four  cases,  e f f e c t s o f t h e t r e a t m e n t s on t h e  eventual  i n s t a r s a r e shown  In  Fecundity  Maximum mean d a i l y  period  varied  f r o m 4.3  i n the JHA-treated  duction dity and  31-33  i n Figs.  rose  sharply  7-10.  24 h r s . o f  o n t h e s e c o n d day  r e a c h e d peak l e v e l s by t h e 5 t h o r 6 t h day when i t s t a b i -  lized.  4.6  assess-  29-32  females produced  l a r v a e were u s u a l l y p r o d u c e d w i t h i n  a d u l t emergence. and  o v e r t h e 10-day  c o n t r o l females produced  o f f s p r i n g whereas k i n o p r e n e - t r e a t e d offspring.  groups  t o 5.6  treated (f)  t o 3.0 a n d 2.6  and more  this,  kinoprene.  t o 3.1  nymphs o f a l a t e  reprofecun-  i n the c o n t r o l  adults  nymphs o f a p t e r o u s  nymphs b o r n t o a l a t e a d u l t s , were s e n s i t i v e to kinoprene.  insects died within avoid  However, a f t e r t h i s ,  so t h a t by t h e 1 0 t h day, mean  Compared t o a p t e r i f o r m  smaller  this  insects, respectively.  Apteriform  apteriform  observed during  i n c o n t r o l s a n d f r o m 4.5 t o  insects.  decreased' slowly  r a n g e d between 2.6  fecundity  adults,  generally  O v e r 9 0% o f t h e  12 h r s . o f k i n o p r e n e - t r e a t m e n t .  To  s u b s e q u e n t e x p e r i m e n t s were p e r f o r m e d w i t h A l l t h e i n s e c t s i n b o t h t h e c o n t r o l and  10  ppm  treated  Table  5.  Overall  effect  fecundity adults (5-6  o f 10 ppm  of apteriform  k i n o p r e n e on t h e nymphs o f  up t o 10 d a y s a f t e r  replicates/treatment;  1  apterous  adult one  emergence  insect/  replicate). Average  fecundity/female/day  Stage t r e a t e d  Control  Kinoprene-treated  1st  instar  3.2±0.0  2nd  instar  3 .2±0.l  3rd  instar  3.2±0.2  a  3.1±0 . 0  4th  instar  3.2±0.0  a  3.2±0.0  3.2±0 . 0 a  3.2±0.1  a  a  a  a  i  1  2  3  \  4 Age  Figure  7.  I  1  !  !  5  6  7  8  !  i  9 1 0  (days)  F e c u n d i t y o f n o r m a l a p h i d s ( •— • ) and t h o s e t r e a t e d w i t h 10 ppm kinoprene ( A ) as f i r s t i n s t a r a p t e r i f o r m l a r v a e o f a p t e r o u s adults.  Fecundity cQ SH fD  to 1—  00  u_i hrj f_) O fD 4 O L-.  < T 3 $_ OJ ^ 3 CD" 3 Q> H-  0  **  OJ  o  KJ  rt  o Hi  3  CD CD O  *  5  n  3  O CD . r_  0)  0) w fr —  Oi I PJ C  M rt cn •  ! U 3^  > LQ CD  •  I I  HC^ cn  Pi OJ  cn  co  o  cn CD  o O  3 ' 0) P 3 3 Ccn rt  rt  3" 0) o 0) T3  rt CD l-i HHi  CO  cn CD  rt H CD  0) rt  O CD  3  (offspring/female/day)  Cb  s: rt  3"  >o  I-  Fecundity  *1  CD  Cn  -r—  l_.l_.hj 0) O fD H O <V C OJ 3 fD 3 f_ I- 1  O *• r t Hi  H-"<  PJ  O o H Hi fD 3 O fD l-i  rt fD H O  co  3  OJ • PJ  i  d i TJ tprt cn cn  fD  OJ  0) cn  1  tn rt  er H-  c_  I  Cn  O  OJ  cn 3CL rt rt H er O OJ cn TJ fD rt fD r t H h H - fD H i 0) O rt H fD B  Ch  : t, •rt er  9£  (offspring/female/day)  CO  NO  1  J  1  !  2  i  3  l  4  Age  i  i  i  I  5  6  7  8  :  i  !  9  10  (days) co  Figure  10.  F e c u n d i t y o f n o r m a l a p h i d (• 10 ppm k i n o p r e n e (A •<->) as larvae of apterous adults.  • ) and t h o s e t r e a t e d w i t h fourth instar apteriform  ^  38  groups developed However, l i k e  i n t o normal-looking apterous  adults.  i n t h e p r e v i o u s e x p e r i m e n t s , m o l t i n g was  d e l a y e d about 1 day i n t r e a t e d f i r s t and second  instar  nymphs a n d 7 o r 8 h r s . i n t h i r d a n d f o u r t h i n s t a r s .  Again,  J H A - t r e a t e d nymphs d i d n o t u n d e r g o any s u p e r n u m e r a r y before molting into  stage  adults.  T a b l e 6 shows t h a t k i n o p r e n e h a d no s i g n i f i c a n t e f f e c t o n r e p r o d u c t i o n . O v e r a 10-day p e r i o d , 2.8 t o 3.2 s p r i n g were p r o d u c e d treated females.  off-  d a i l y by t h e c o n t r o l and k i n o p r e n e -  I n a l l instars., l a r v i p o s i t i o n began  2 4 h r s . o f a d u l t emergence  (Figs. 11-14).  within  Reproduction  r a t e s r o s e r a p i d l y and r e a c h e d a peak by t h e 5 t h o r 6 t h day. D u r i n g t h i s p e r i o d , maximum mean d a i l y  fecundity  varied  f r o m 4.0 t o 4.8 i n t h e c o n t r o l s , a n d f r o m 4.0 t o 4.4 i n t h e treated insects.  After this,  r e p r o d u c t i o n d e c r e a s e d so  t h a t b y t h e 1 0 t h d a y , mean f e c u n d i t y r a n g e d b e t w e e n 2.6 t o 3.0 a n d 2.6 - 3.2 i n c o n t r o l a n d t r e a t e d i n s e c t s , (g)  Apterous adults  respectively,  (apterae-producers)  U n l i k e a p t e r i f o r m nymphs, n e w l y e c d y s e d  apterous  a d u l t s w e r e l e s s s e n s i t i v e t o 65 ppm k i n o p r e n e a s o n l y 1 0 % m o r t a l i t y was o b s e r v e d .  As i n p r e v i o u s s t u d i e s ,  kinoprene  h a d no s i g n i f i c a n t e f f e c t o n r e p r o d u c t i o n ( F i g . 1 5 ) . I n b o t h c o n t r o l a n d k i n o p r e n e - t r e a t e d f e m a l e s , maximum f e c u n d i t y o c c u r r e d on t h e 6 t h day.  However, a f t e r  daily this,  Table  6.  O v e r a l l e f f e c t o f 10 ppm fecundity adults (5-6  of apteriform  k i n o p r e n e on t h e nymphs o f  alate  (up t o 10 d a y s a f t e r a d u l t  r e p l i c a t e s / t r e a t m e n t , one  emergence  insect/  replicate). Average  fecundity/female/day  Stage t r e a t e d  Control  1st  instar  2.9±0.1  C  2.9±0.1  2nd  instar  3.2±0.1  J  3.0±0.1*  3rd  instar  2.8±0.1  J  3.0±0  4th  instar  3.0±0  a  Kinoprene-treated  a  3.0±0.3  £  Fecundity  (offspring/female/day)  ffD  O H  ^ O  fD  hh O  0) pj rt (D 0) Ch  CO  'O 3  W rt 3 O o c •« hh H H 3 r t fD O cn 3 . fD 1  I I I i •  &) *v  co  tr.  >  Oi cn  CD  H-  Oi P> ^< cn  P> cn hh  Oi  o  Hhi  cn  rt — P)  3  cn Oi  rt  rt 3  J  hi  O pj cn  ^ fD rt CD r t  CO  hi  CD p> rt CD Oi i— s: pj p. hi r t < 3*  >o  1  P)  CD  I — h  Fecundity d H  ro  O M Hi O  ^ fD O P> 13 d M 13 3 pj g rt (D W rt  Cu d M rt CD •  O 13 H fD 3 fD  ho  O Hi  CO  3 O H  3  f |  ! ! ^  » Hcn  PJ — . cn • cn fD  > CD  PJ cn  o  o  O 3 Ch  H3 cn rt PJ H  PJ 3 Ch  rt 3" PJ O 13 cn rt fD fD i-i rt H Hi fD O P) h rt 3 CD Ch Hrt  tr  (offspring/female/day)  CO  •o  CO  On  Fecundity  4  (offspring/female/day)  HiQ  0  H CD  to  M  to  0 I— •Hi O  1  r-"Tj OJ g r+  hrj  -fr-e—H  CD Oc  O-  CD ;v r t  rO  c_ o o  r_ TS H i H H r t CD 3 01 3 O • CD H  CO  3  —  0)  >  J OJ  I.  I 0)  cQ CD  H-  cn  cn  Pi  0) cn  rt  o  cr Hi-i  Pi  H- 0)  3cn rt 0) H  Pi  rt cr  O  0) cn  rt  CD  CD r t H H H- CD H i 0)  O rt i-i  3  CD  Pi  3  H-  rt  cr  3fr  CO  to  Oi  —I  1  2  3  4  5 Age  Figure  14.  6  7  8  9  10  (days)  F e c u n d i t y o f n o r m a l a p h i d s (• • ) and t h o s e t r e a t e d 10 ppm k i n o p r e n e (_) as f o u r t h i n s t a r a p t e r i f o r m of a l a t e adults.  with larvae  T  J  1  2  3  !  4  5 Age  Figure  15.  I  6  L  7  8  9  10  (days)  F e c u n d i t y o f n o r m a l a p h i d s (•; •) and t h o s e t r e a t e d w i t h 65 k i n o p r e n e (* --*) a s n e w l y - e c d y s e d a p t e r o u s a d u l t s .  ppm  45  fecundity declined  a b r u p t l y so t h a t b y t h e e n d o f 1 0 t h d a y ,  mean f e c u n d i t y was 3.2 i n t h e t r e a t e d the  a n d 3.3 i n  controls. (h)  Apterous To  adults  within  (alate-producers)  a s s e s s t h e p r e n a t a l e f f e c t s o f JH on w i n g d e -  velopment, kinoprene  (65 ppm) was a p p l i e d  12 h r s . o f t h e i r e m e r g e n c e .  r e a r e d t o g e t h e r i n each for  insects  cage.  t o apterous  Twenty f e m a l e s  adults  were  O f f s p r i n g were c o l l e c t e d  daily  3 d a y s a n d r e a r e d u n d e r crowded c o n d i t i o n s t o f a v o u r  the p r o d u c t i o n o f a l a t i f o r m s . the o f f s p r i n g 76.6%  developed  of the offspring  developed  into  born  apterae  o n l y 10% o f  (Table 7 ) .  In contrast,  to kinoprene-treated insects  normal-looking apterous  t r e a t m e n t h a d no e f f e c t spring  into  In t h e c o n t r o l s ,  adults.  on d e v e l o p m e n t a l  The k i n o p r e n e -  times o f the o f f -  a s t h e y emerged as a d u l t s a t t h e same t i m e  as the "  controls.  B.  COMBINED EFFECTS OF PRECOCENE AND (a)  First  and s e c o n d  instar  KINOPRENE  a p t e r i f o r m nymphs o f a p t e r o u s  adults When 0.5 ug p r e c o c e n e - I I was a p p l i e d first no  and s e c o n d  significant  instar  lethal  was d e l a y e d a t e a c h molted  about  topically to  a p t e r i f o r m nymphs o f a p t e r o u s  e f f e c t s were a p p a r e n t .  subsequent  adults,  However, e c d y s i s  nymphal s t a g e a s c o n t r o l  1 t o lh d a y s s o o n e r  aphids  than t h e p r e c o c e n e - t r e a t e d  T a b l e 7.  A p t e r a e p r o d u c t i o n by a p t e r o u s f e m a l e s w h i c h were t r e a t e d w i t h k i n o p r e n e w i t h i n 12 h r s . o f a d u l t  Treatment  Control  Age o f m o t h e r s when l a r v a e collected  No. o f l a r v a e collected  65 ppm  emergence. No. o f l a r v a e molting into apterous adults  % apterous a d u l t s produced  1-day-old  40  5  12.5  2  "  40  3  7.5  3  "  40  4  10. 0 ±=10.0  Kinoprene  1-day-old  40  30  75. 0  2  "  40  32  80.0  3  "  40  30  75. 0 ±=76.6  CTl  47  insects.  Although a l l the  normal-looking  apterae,  lower than i n the days of a d u l t of  3.2  was  the  offspring/day  produced only  their  controls  life,  0.7  l a t t e r i n s e c t s developed f e c u n d i t y was  (Table  8).  During the  whereas the p r e c o c e n e - t r e a t e d  increased  the  controls  II  and  k i n o p r e n e on  The  8).  The  fecundity  are  never reached l e v e l s  s t i m u l a t o r y e f f e c t s o f k i n o p r e n e on f e m a l e s were n o t  a f t e r t h e JHA  been a p p l i e d .  f e c u n d i t y was (b)  T h i r d and apterous  A l t h o u g h no  apparent u n t i l  Following  17.  of 1 o r 2 days  this,  the  2 offspring/day.  By  kinoprenethe  fourth instar apteriform  nymphs o f  10th  adults  s i g n i f i c a n t l e t h a l e f f e c t s were o b s e r v e d a p p l i e d t o t h i r d and was  veloped i n t o normal-looking  and  A l l the  apterae.  lower than the first  fourth instar  delayed f o r about 7 or  a t each subsequent nymphal m o l t .  a 10-day p e r i o d ,  fecundity  and  controls,  a p t e r i f o r m nymphs, e c d y s i s  precocene-treated  precocene-  comparable to t h a t of the  when p r e c o c e n e - I I was  significantly  females  shown i n F i g s . 16  precocene-treated  day,  average  emergence,  d a i l y e f f e c t s of  t r e a t e d females produced n e a r l y  10  However, when k i n o p r e n e  s i g n i f i c a n t l y but  (Table  had  first  c o n t r o l f e m a l e s p r o d u c e d an  offspring/day.  in  was  significantly  a p p l i e d t o these i n s e c t s 4 days a f t e r a d u l t  fecundity  into  8 hrs.  t r e a t e d nymphs  However, t h e i r  c o n t r o l s but  higher  fecundity than  second i n s t a r s ( T a b l e 8).  f e m a l e s i n c o n t r o l and  de-  t r e a t e d groups  Over pro-  T a b l e 8.  E f f e c t s o f t o p i c a l a p p l i c a t i o n o f p r e c o c e n e - I I on t h e f e c u n d i t y o f a p t e r i f o r m nymphs o f a p t e r o u s a d u l t s .  E x c e p t on a p t e r o u s  adults,  k i n o p r e n e was a p p l i e d t o t h e p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t e m e r g e n c e (5-6 r e p l i c a t e s / t r e a t m e n t , Average Stage o f precocene application  Control  fecundity/surviving  Precocene-treated  1st  instar  3, 2±0.0  2nd  instar  3, 2±0.1'  0.7-iO.O  3rd  instar  3, 2±0.1  4th  instar  3, 2±0. 0  Apterous  adults  3, 2+0.0  C  £  e  £  one i n s e c t / r e p l i c a t e ) * .  0.7±0.0  female/day Precocene-kinoprene treated  1.2±0.1 1.4±0.0  (  1.8±0.2*  2.3+0.1  C  2.5±0.0  2.7 + 0.0*  1  1  0.6±0.1*  1.7±0. 0 d**. 2.5±0.0 C  * Mean d i f f e r e n c e s a t 5% l e v e l u s i n g Duncan's M u l t i p l e Range t e s t * * K i n o p r e n e a p p l i e d 24 h r s . a f t e r a d u l t e m e r g e n c e .  CO  5  1 F i g u r e 16.  2  3  4  Age  5 (days)  6  7  8  9  F e c u n d i t y o f c o n t r o l a p h i d s (• • ) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I (--) a s f i r s t i n s t a r a p t e r i f o r m nymphs o f a p t e r o u s a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e ( o — — — — — o ) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  Age F i g u r e 17.  (days)  F e c u n d i t y o f c o n t r o l a p h i d s (• • ) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I (-) a s s e c o n d i n s t a r a p t e r i f o r m nymphs o f a p t e r o u s a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e (o o) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  51  duced  a n a v e r a g e o f 3.2 a n d 1.5-2.5 o f f s p r i n g / d a y ,  respectively.  K i n o p r e n e a p p l i c a t i o n t o p r e c o c e n e - t r e a t e d i n s e c t s on t h e i r 4th day o f a d u l t l i f e  increased  f e c u n d i t y o f emergent t h i r d  fecundity  significantly.  i n s t a r s was s i g n i f i c a n t l y  The  lower  t h a n i n t h e c o n t r o l s whereas t h e f e c u n d i t y o f t h e emergent f o u r t h i n s t a r s was c o m p a r a b l e  to the controls  ( T a b l e 8 ) . The  d a i l y e f f e c t s o f p r e c o c e n e - I I and k i n o p r e n e on t h e f e c u n d i t y o f t h e t w o i n s t a r s a r e shown i n F i g s . 1 8 - 1 9 . p l i c a t i o n r e s u l t e d i n an immediate  increase  Kinoprene api n fecundity of  both i n s t a r s . (c)  Apterous  adults  When 0.2 u g p r e c o c e n e - I I was a p p l i e d t o n e w l y e c d y s e d a p t e r o u s a d u l t s , f e c u n d i t y was s i g n i f i c a n t l y (Table 8 ) .  Compared t o t h e c o n t r o l  an a v e r a g e o f 3.2 o f f s p r i n g / d a y , females produced  females, which  When k i n o p r e n e was  a p p l i e d t o t h e s e i n s e c t s 24 h r s . a f t e r t h e i r a d u l t  Kinoprene a p p l i c a t i o n t o 4-day-old  treated adults also contributed increase  fecundity  s o t h a t b y t h e 7 t h d a y , i t was c o m p a r a b l e  controls.  i n fecundity.  emergence,  were n o t a p p a r e n t u n t i l  l a t e r , ( F i g . 2 0 ) . However, a f t e r t h i s , sharply  produced  the precocene-treated  0.8 o f f s p r i n g d a i l y .  t h e e f f e c t s on r e p r o d u c t i o n  reduced  3 days  increased to the  precocene-  t o a d e l a y e d b u t pronounced  1  2  3  4  5  Age F i g u r e 18.  6  7  8  9  10  (days)  F e c u n d i t y o f c o n t r o l a p h i d s (• -•) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I (-*) a s t h i r d i n s t a r a p t e r i f o r m nymphs o f a p t e r o u s a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e (o o) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  1  j  I  2  3  I  1  4  5  Age g u r e 19.  1 6  1 7  1 8  ! 9  1 10  (days)  F e c u n d i t y o f c o n t r o l a p h i d s (• e) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I A) as f o u r t h i n s t a r a p t e r i f o r m nymphs o f a p t e r o u s a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e (o o) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o indicated.  Fecundity  (offspring/female/day)  PIO  d H CD  -r—  O  rt  3  OJ rt  •CS CD  Ch  CD  O O o  prt  OJ CD H i 13 r t CD CD I H rt H. OJ CD Ch 0) d rt M CD rt  OJ  CD  CD  ch  3  CD  H -1 CD 3 O CD  CD O d 3  P-  d  PJ_)  d M13 rt cn CD  hi o o • o 3 t-3  ty  O O. CD  3 CD  rt H O  0)  I Ti  cn ro O rt cn  CD H 3" HI-H HI p-  d-  O Hi  |  i-i  o\  \  cn  CO  O Hi  CD  PJ  —  cn  O lien  -1 CD  —  Ch 0J  K:  cn  ui 13 .13 OJ cn  H cn O  3  *T  P3  P3 Ch O p13 OJ — - P, rt CD CD O 3 Ch hj CD  o  OJ 13  £—13  OJ ^ cn  H p.  CD  Ch  3 CD  0) 3 C-  03  £ M rt ^  I  CD  O  Ch  tr o cn CD  cn  rt H  CD  CD  S—>  OJ rt CD  Ch  O  L  CO  Ol  55  (d)  Apteriform  nymphs o f a l a t e a d u l t s  Experiments performed with precocene-II and k i n o prene on the f i r s t and second i n s t a r a p t e r i f o r m nymphs o f a l a t e a d u l t s produced more o r l e s s the same r e s u l t s as w i t h a p t e r i f o r m nymphs o f apterous a d u l t s . ference  No s i g n i f i c a n t  dif-  i n l e t h a l i t y was observed between the t r e a t e d and  control insects.  However, e c d y s i s was again delayed a t each  subsequent nymphal stage i n a l l the t r e a t e d i n s e c t s . A l though a l l the precocene-treated  i n s e c t s developed i n t o normal  l o o k i n g apterae, t h e i r f e c u n d i t y was s i g n i f i c a n t l y compared to the c o n t r o l s observed during  (Table 9 ) . Mean d a i l y  reduced  fecundity  the f i r s t 10 days o f a d u l t l i f e ranged from  2.2 to 3.0 i n the c o n t r o l s and from 0.8 t o 2.0 i n the t r e a t e d aphids.  When kinoprene was a p p l i e d t o these i n s e c t s 4 days  a f t e r t h e i r a d u l t emergence, f e c u n d i t y i n c r e a s e d  significantly.  However, f e c u n d i t y never reached the l e v e l s o f c o n t r o l s i n the emergent JHA-treated f i r s t and second i n s t a r s ( F i g s . 2122).  In t r e a t e d t h i r d and f o u r t h i n s t a r s , i t was  equivalent  to t h a t o f c o n t r o l s by the 5th-7th day o f a d u l t emergence ( F i g s . 23-24).  T a b l e 9.  E f f e c t s o f t o p i c a l a p p l i c a t i o n o f p r e c o c e n e - I I on t h e f e c u n d i t y o f a p t e r i f o r m nymphs o f a l a t e a d u l t s . 4? p r e c o c e n e  - t r e a t e  d  to the  i n s e c t s f o u r d a y s a f t e r a d u l t emergence (5-6  replicates/treatment,  one i n s e c t / r e p l i c a t e ) *  Average Stage o f precocene application  K i n o p r e n e was a p p l i e d  Control  fecundity/surviving  Precocene-treated  female/day Precocene-kinoprene treated  1st i n s t a r  2.7±0.0  a  0.8±0.0  b  1.1±0.0  C  2nd  instar  3.0±0.1  a  0.8+0.1  b  1.3±0.0  C  3rd  instar  2.8±0.1  a  1.5±0.2  b  2.6+0.0  a  4th  instar  3.0+0.1  a  2.0±0.7  b  1.5±0.1  C  * Mean d i f f e r e n c e s  a t 5% l e v e l u s i n g Duncan's M u l t i p l e Range t e s t  5  1  2  3  4  5  Age F i g u r e 21.  6  7  8  9  10  (days)  F e c u n d i t y o f c o n t r o l a p h i d s (• •) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I (* ±) as f i r s t i n s t a r nymphs o f a l a t e a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e (o o) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  5T  Age F i g u r e 22.  (days)  F e c u n d i t y o f c o n t r o l a p h i d s (•-———•) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I *•) as. s e c o n d i n s t a r a p t e r i f o r m nymphs o f a l a t e a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e (o o) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  co  5  Age Figure  23.  6  8  (days)  F e c u n d i t y o f c o n t r o l a p h i d s (• •) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I (* *) as t h i r d i n s t a r a p t e r i f o r m nymphs o f a l a t e a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e (o o) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  CD  4  5  Age F i g u r e 24.  6  8  10  (days)  F e c u n d i t y o f c o n t r o l a p h i d s (• •) and t h o s e t r e a t e d w i t h 0.5 ug p r e c o c e n e - I I --) as f o u r t h i n s t a r a p t e r i f o r m nymphs o f a l a t e a d u l t s . The e f f e c t s o f 10 ppm k i n o p r e n e (o o) a p p l i e d t o p r e c o c e n e - t r e a t e d i n s e c t s f o u r d a y s a f t e r a d u l t emergence a r e a l s o i n d i c a t e d .  cn o  61  DISCUSSION  A.  POLYMORPHISM IN THE  GREEN PEACH APHID  Although polymorphism i n f l u e n c e d by Hille  a variety  i n aphids  of environmental  R i s Lambers 1966),  i s known t o stimuli  be  (Lees  1966,  t h e p h y s i o l o g i c a l mechanisms w h i c h  c o n t r o l morph d e t e r m i n a t i o n a r e p o o r l y u n d e r s t o o d .  In  current  to long-  day  study,  c o l o n i e s o f M^  p e r s i c a e were e x p o s e d  c o n d i t i o n s to promote the e x c l u s i v e  production of  v i r g i n o p a r a e w h i c h a r e p a r t h e n o g e n e t i c and addition,  a d u l t v i r g i n o p a r a e and  under e i t h e r  isolated  their  the  adult  viviparous.  In  o f f s p r i n g were r e a r e d  c o n d i t i o n s to favour the p r o d u c t i o n of  a p t e r o u s morphs o r crowded c o n d i t i o n s t o p r o m o t e  formation  of  of  alates.  In the l a t t e r  i n s t a n c e , the response was  n o t a b s o l u t e as n e a r l y  apterous  virginoparae  to crowding  the progeny,  w h i c h were a l s o m a i n t a i n e d u n d e r crowded into apterae.  con-  developed  crowding  has b e e n shown t o f a v o u r a l a t e p r o d u c t i o n , v a r i a b l e  However, i n t h i s to  crowding  this  where  ( L e e s 1967,  maternal  1980).  s p e c i e s , many a p t e r o u s v i r g i n o p a r a e r e s p o n d  by p r o d u c i n g e x c l u s i v e l y  s u g g e s t s t h a t M^  t h a n M^_ p e r s i c a e , to  are a l s o produced  viciae,  of  ditions,  proportions of alates  I n M^  10%  viciae  alate offspring.  i s more s e n s i t i v e  somewhat d i f f e r e n t  s i m u l a t e crowded c o n d i t i o n s .  to  Although  crowding  t e c h n i q u e s were  employed  In the p r e s e n t study,  these  62  conditions  were c r e a t e d  parae i n t o c l i p - c a g e s whereas, i n the  by  t r a n s f e r r i n g 25 a p t e r o u s v i r g i n o -  attached  t o C h i n e s e cabbage p l a n t s ;  i n v e s t i g a t i o n s of Lees  (1967, 1 9 8 0 ) ,  10  a d u l t s were t r a n s f e r r e d i n t o specimen t u b e s , crowded f o r  24  hrs.,  fre-  then returned  t o bean p l a n t s .  quency of a l a t e f o r m a t i o n not  only  crowding but In  be  in  v i c i a e may  a l s o temporary  persicae,  be  a t t r i b u t e d to  starvation.  several morphological c r i t e r i a  used t o d i s t i n g u i s h between the  These d i f f e r e n c e s  Therefore, the high  a l a t e and  can  a p t e r o u s morphs.  f i r s t become a p p a r e n t i n t h e  third  instar  when w i n g b u d s become v i s i b l e e x t e r n a l l y i n p r e s u m p t i v e a l a t e s . The  a p t e r o u s nymphs a r e  and  lack swellings  development of c a u d a and  an  i n the  with  the  Except for a  pointed  r e t a i n many n y m p h a l c h a r a c t e r i s t i c s .  unpigmented body, reduced t h o r a c i c  example, the  an  sclerotizationandthe  appear t o complement w i n g  thorax.  important adaptation  bonds between N - a c e t y l  d o p a m i n e and  heavily  This s c l e r o t i z a t i o n  to f l i g h t .  z a t i o n p r o c e s s i s known t o i n v o l v e t h e  very  development.  c u t i c l e becomes d a r k l y p i g m e n t e d and  s c l e r o t i z e d p a r t i c u l a r l y i n the  little These  I n c o n t r a s t , metamorphic changes are  p r o n o u n c e d i n a l a t e s and  represents  i n appearance  e n l a r g e d g e n i t a l p l a t e , a d u l t apterae undergo  absence of o c e l l i .  For  prothorax associated  f l i g h t muscles i n a l a t e s .  m e t a m o r p h o s i s and include  somewhat more g l o b u l a r  The  formation  proteins  of  i n the  scleroticovalent exocuticle  63  (Chapman 1 9 7 5 ) . stronger  and  muscles. present  a result,  better able  Ocelli  and  p h o t o and  the  thoracic cuticle  i s rendered  to support c o n t r a c t i o n s of the  secondary antennal  i n adult alates.  i s debatable,  s e n s o r i a are  Although t h e i r  c h e m o r e c e p t o r s and host-plant  may  be  also  importance i n  important  in  l o c a t i o n (Wellington  flight as  flight  1974;  Goodman  B r o m l e y e t a l . 1979) . I n m o s t h e m i m e t a b o l o u s and  paurometabolous i n s e c t s ,  n y m p h a l d e v e l o p m e n t , m e t a m o r p h o s i s and separated  temporally  (Chapman 1 9 7 5 ) .  reproduction  l a r v i p o s i t i o n u s u a l l y b e g i n s on  day  As  of adult l i f e .  ment and  i t i s inconceivable  e m b r y o g e n e s i s c o u l d o c c u r i n one  that reproductive Although the  the  that oocyte day,  first develop-  stages.  s i m u l t a n e o u s d e v e l o p m e n t o f t h e m o t h e r and the generation  t i m e and  i n c r e a s e s , t h e phenomenon p r e s e n t s  p h y s i o l o g i s t s t o e x p l a i n how  of the o f f s p r i n g d i f f e r s  favour  from t h a t of the  her rapid  a challenge  such development i s  s i t u a t i o n becomes e v e n more c o m p l i c a t e d  and  i t i s apparent  development b e g i n s i n the nymphal  o f f s p r i n g would shorten population  are  However, i n a p t e r o u s  alate virginoparae,  The  flight  these sensory organs are b e l i e v e d to serve  n a v i g a t i o n and 1975;  As  coordinated.  when t h e  mother.  to  morphology  64  B.  ROLE OF As  believe and  JUVENILE HORMONE described  t h a t JH  plays  polymorphism.  histological  Elliott  and  postulated  JH  that  the  r o l e i n aphid  corpora  morphogenesis  s o l e l y on  allata  (CA),  the  retention of  i n h i b i t e d wing development. titers  declined  juvenile They  T h i s w o u l d e x p l a i n why  changes i n the  CA  of glandular  JH  Moreover,  synthesis.  are  growth-regulating  o f JHA  in various  aphid  provide  of  the  role of  i n aphid  an  attempt to r e s o l v e  JH,  the  no  controversy  stages of  choice  biologically the  a n a l o g u e has  (Lees 1980).  JHI,  properties  surrounding the topically  has  proven  r e c e n t l y b e e n shown t o be  a c t i v e i n a p h i d s as hormone  JHA  numerous  reproduction.  a p t e r o u s morphs o f  of kinoprene over other  because the  form o f  a l a t e and  and  clear indication  m o r p h o g e n e s i s and  k i n o p r e n e a n d / o r p r e c o c e n e were a p p l i e d  various The  JH  alates  not i n d i c a t i v e  s t u d i e s , w h i c h have e x a m i n e d t h e species,  but  these hypotheses  suggested t h a t volumetric and  also  The u l t r a s t r u c t u r a l  (1976) c h a l l e n g e d  activity  l e v e l s of  i n apterous adults  produce apterous o f f s p r i n g .  of Leckstein  the  White  (1975) s u g g e s t e d t h a t h i g h  in alate adults.  invariably  some i n v e s t i g a t o r s  conclusions  nymphs f a v o u r e d  characteristics  studies  important  appearance of  i n apteriform  increased  an  Introduction,  Basing t h e i r  (1965, 1968b) and JH  i n the  (JH)  In  role to  of  the  persicae. advantageous as  a naturally-occurring  F u r t h e r m o r e , as w i l l  become  65  evident study  s h o r t l y , the  a p p l i c a t i o n methods e m p l o y e d  have a d v a n t a g e s o v e r  treated  foliage  r o l e o f JH (a)  other bioassay  or a r t i f i c i a l  i n the  individual  methods  stages  can  be  insects, allatectomy  o v a r i a n development  1976).  this  Since  i n h i b i t i o n can  (Engelmann 1970; be  r e v e r s e d by  i t has  b e e n e s t a b l i s h e d t h a t JH  deposition  i n the  developing  role  in reproduction  o f JH  reproductive f o r 48  the  h r s . was  direct  or  i s not  to the  Mittler  0.1%  species, In A^  different  has  JHA-treated from  that  no  influence  are  unclear  adults,  because  a l s o h i g h l y r e p e l l e n t and  fabae  kinoprene.  the  fabae,  f a c t o r i n apterous  e t a l . (1976) r e a r e d  o b s e r v a t i o n p e r i o d , the t o t a l t r e a t e d and  was  JHA  yolk  (Hangartner e t a l . 1971).  v i r g i n o p a r a e o f A^  plants treated with  topical  ( H a n g a r t n e r e t ajL. 1971) .  reproduction  study  female aphids  techniques,  g y n o p a r a e and  the  foliage  a limiting  e f f e c t s o f JH on  Gilbert  a d u l t s r e a r e d on  significantly  untreated  the analogue used i n t h i s  similar  In a p h i d  remains u n c e r t a i n .  not  treat-  stimulates  t h i s w o u l d a p p e a r t o i n d i c a t e t h a t JH  reproduction  toxic  oocytes.  r a t e of apterous  o f a d u l t s r e a r e d on  on  JHA-  evaluated,  or precocene  treatment,  Although  (ie.  d i e t s ) i n t h a t the p o t e n t i a l  ment i n h i b i t s  plants  this  Reproduction In other  the  in  and  During  fourth  persicae the  Using instar on  three-week  number o f p r o g e n y p r o d u c e d  c o n t r o l groups d i d not  differ  greatly.  by However,  66  during  the f i r s t  initiated  few d a y s o f a d u l t  sooner  and o c c u r r e d  kinoprene-treated (1976) and  suggested  that  larviposition  at a faster rate  t h a t J H may i n both  CA a c t i v i t y  stimulate  species.  Mittler  ovarian  also  of c e r t a i n alate c h a r a c t e r i s t i c s , notably  muscles.  This  Furthermore,  slow r e p r o d u c t i v e  Mittler  nymphs t h a n t h o s e o f  e t a_l. s u g g e s t e d  o f l e s s J H o r g o n a d o t r o p i c hormone.  c o m p a r a b l e g o n a d o t r o p i c r o l e o f t h e CA was n o t (MacKauer e t a _ l . 1979) .  a p p l i c a t i o n o f 1.0 ug p r e c o c e n e - I I 3- t o 5 - h r . - o l d ty.  Although  offspring were  In t h i s  a small  A  demonstrated  species,  topical  ( d i s s o l v e d i n acetone)  apterous virginoparae  produced  that the  m a t u r a t i o n o f nymphal g y n o p a r a e may r e s u l t  from t h e p r o d u c t i o n  i n A^ pisum  the f l i g h t  i n h i b i t i o n w o u l d e x p l a i n why more embryos  i n the o v a r i o l e s o f apteriform  alatiforms.  specu-  i n h i b i t the  formation  develop  et a l .  development  The a u t h o r s a l s o  i n t h e nymphs m i g h t  was  i n the  a p h i d s t h a n .in t h e c o n t r o l s .  embryogenesis  lated  life,  to  r e s u l t e d i n 63% m o r t a l i -  d e c l i n e was o b s e r v e d  i n t h e number o f  by t h e s u r v i v o r s , no s t e r i l a n t  effects  apparent. I n M^ p e r s i c a e ,  precocene-II within  topical  ( d i s s o l v e d i n DMSO) t o a p t e r o u s  2 to 3 hrs. of t h e i r  m o r t a l i t y but induced almost female  a p p l i c a t i o n o f 0.5 ug  produced  only  virginoparae  emergence r e s u l t e d i n no complete  one o f f s p r i n g / d a y  sterility.  adult  Each  b u t t h e nymphs  treated died  67  w i t h i n 24 h r s . o f b i r t h . effects,  t h e precocene  In order t o avoid these  d o s a g e was r e d u c e d t o 0.2  A t t h i s dosage, l a r v i p o s i t i o n d e c l i n e d almost One d a y a f t e r a p p l i c a t i o n , produced  toxic ug/aphid.  immediately.  t h e precocene-treated females  o n l y one o f f s p r i n g / d a y w h e r e a s t h e c o n t r o l s  duced t h r e e t o f o u r o f f s p r i n g d a i l y .  Reduced  pro-  larviposition  i n t h e p r e c o c e n e - t r e a t e d f e m a l e s c o n t i n u e d f o r a t l e a s t 10 days.  However, t h e i n h i b i t o r y e f f e c t s o f p r e c o c e n e  were  r e v e r s e d when t h e f e m a l e s w e r e t r e a t e d w i t h 65 ppm k i n o p r e n e . K i n o p r e n e was more s t i m u l a t o r y t o l a r v i p o s i t i o n when i t a p p l i e d w i t h i n 24 h r s . o f t h e p r e c o c e n e i t was a p p l i e d  t r e a t m e n t t h a n when  4 days a f t e r t h e precocene  treatment.  In  b o t h i n s t a n c e s , a n i n c r e a s e i n l a r v i p o s i t i o n was n o t e v i d e n t f o r a t l e a s t 2 d a y s a f t e r k i n o p r e n e was a p p l i e d .  Comparable  e f f e c t s w e r e o b s e r v e d when 0.5 u g p r e c o c e n e - I I was a p p l i e d t o f i r s t and second  i n s t a r a p t e r i f o r m nymphs o f a p t e r o u s o r  alate virginoparae. they produced  A f t e r t h e nymphs h a d e m e r g e d a s a d u l t s ,  l e s s t h a n one o f f p s p r i n g / d a y b u t when  treated  w i t h kinoprene, l a r v i p o s i t i o n r a t e s doubled a f t e r 1 o r 2 days. In c o n t r a s t , precocene  t r e a t m e n t s a p p l i e d t o t h i r d and f o u r t h  i n s t a r a p t e r i f o r m nymphs o f a p t e r o u s o r a l a t e a d u l t s w e r e less  inhibitory.  significantly  Although the treated insects  fewer o f f s p r i n g than t h e c o n t r o l s ,  produced larviposition  r a t e s w e r e two o r t h r e e t i m e s h i g h e r t h a n t h o s e o f a d u l t s  68  w h i c h had  been t r e a t e d w i t h  i n s t a r nymphs. w h i c h had instars to  been t r e a t e d w i t h  aphids.  role  attain  in  a l l stages.  (Unnithan  and  an  However, t h e to  Nair  stage.  If this  of t h i r d  and  presented  Nair  adults  fourth  comparable  final To  effect  particularly  pattern of Moulting  of  CA the  must be  active  appears  i n other  species  nymphal i n s t a r was  account  for this  of  first  to  the  low  sensi-  precocene  o n l y when t h e in  are  CA  are  persicae,  the  apteriforms  are  and  second  instars  Although a d d i t i o n a l evidence  these CA  CA  peach  CA  fourth instar  active.  to support  the  i s tenable  plays  a p t e r o u s morphs  (1979) s u g g e s t e d t h a t  hypothesis  very  i f the  observed  1979), the  i n d i c a t e t h a t the  different (b)  and  i n apterous green  activity  i n a c t i v e whereas the  adults are  studies  second  d e m o n s t r a t e t h a t JH  levels,  As  anti-allatotropic  relatively  be  in reproduction  U n n i t h a n and  active.  and  results  s e n s i t i v e to precocene.  exerts  CA  the  normal r e p r o d u c t i v e  from stage  tivity,  third  r a t e s to l e v e l s  They a l s o i n d i c a t e t h a t  to  least  or  of the c o n t r o l s .  important  vary  p r e c o c e n e as  larviposition  Collectively, an  first  In a d d i t i o n , k i n o p r e n e a p p l i c a t i o n to  increased  those  precocene as  conclusions,  the  will  subsequent  i n a l a t i f o r m nymphs e x h i b i t a  activity.  and  metamorphosis  In e x o p t e r y g o t e s ,  progressive  morphological  changes  69  which occur d u r i n g m o u l t i n g hormonally  (Wigglesworth  and m e t a m o r p h o s i s a r e  1970;  Chapman 1975;  mediated  Gilbert  I n c o n j u n c t i o n w i t h t h e m o u l t i n g hormone, e c d y s o n e , b e e n shown t o programme t h e e p i d e r m a l type of c u t i c l e  ment o f JH  i n t h e programming o f t h e e p i d e r m a l  techniques. performed  by  a variety  on p e n u l t i m a t e  characteristics. administered  o r young nymphal  t o nymphal  i s applied,  characteristics.  h y p o t h e s i z e d t h a t when JH each m o u l t i n g form.  cycle,  the  However, when JH  the i n s e c t  and  develops  the occurrence tous d e c l i n e As  i n s e c t s may From t h e s e  titer  insect  exhibit studies,  develops  d u r i n g the  t h a n a p t e r o u s morphs.  adult  treatments metamorphosis.  i t has  an  of  immature  Thus, i n normal  final  and/  been  initiation  into  when  juvenile  during the c r i t i c a l  an a d u l t .  discussed previously  e x h i b i t e d by most i n s e c t  induced  i t s d o s a g e and  o f metamorphosis i s a t t r i b u t e d  i n JH  treatments  instars  i s p r e s e n t a t the  i s absent  into  has  chemical  stages delay or prevent  the t r e a t e d  involve-  cells  i m p l a n t s o r JHA  o f t h e JHA,  has a  premature development o f  C o n v e r s e l y , CA  D e p e n d i n g upon t h e a c t i v i t y  or adult  The  F o r example, a l l a t e c t o m y o r p r e c o c e n e  p r e c o c i o u s m e t a m o r p h o s i s and  it  t o each moult.  of surgical  JH  to produce  particular  been d e m o n s t r a t e d  prior  cells  1975) .  to a  period,  insects, precipi-  instar.  the developmental  pattern  s p e c i e s i s more e v i d e n t i n a l a t e  I f t h e h o r m o n a l mechanisms  controlling  70  m o u l t i n g and metamorphosis i n a p h i d s a r e s i m i l a r t o t h a t o f other exopterygotes, apteriform by  nymphs s h o u l d be  h i g h e r J H l e v e l s t h a n a l a t i f o r m nymphs.  many a p h i d s p e c i e s , 1971;  characterized  As r e p o r t e d i n  (Lees 1966, 1977, 1980; H a n g a r t n e r e t a l .  H r d y 1974; C l o u t i e r a n d P e r r o n 1 9 7 5 ; M i t t l e r e t a l .  1976) , a l a t i f o r m nymphs o f M_^_ p e r s i c a e  treated with  develop i n t o l a r v a l - a d u l t intermediates. juvenilization exhibited  most J H A - s e n s i t i v e ( L e e s 19 8 0 ) ,  The d e g r e e o f  by t h e emergent a d u l t  when t h e k i n o p r e n e was a p p l i e d .  Unlike  depended upon  M^ v i c i a e w h e r e t h e  s t a g e was t h e t h i r d . i n s t a r a l a t i f o r m nymph  t h e development o f f i r s t and second'  instar alatiforms  o f ML_ p e r s i c a e  was d r a s t i c a l l y a l t e r e d b y  t o p i c a l a p p l i c a t i o n s o f 65 ppm k i n o p r e n e . treated  O v e r 80% o f t h e  i n s e c t s underwent a supernumerary moult b e f o r e de-  veloping  into adults.  A p a r t from t h e presence o f s m a l l  buds, t h e a d u l t s were a l m o s t i n d i s t i n g u i s h a b l e from adults. had  JHA  Furthermore, the i n s e c t s  lacked  an u n d e r d e v e l o p e d g e n i t a l p l a t e .  a pointed  wing  apterous cauda and  As both f e a t u r e s a r e  found i n a p t e r o u s and a l a t e v i r g i n o p a r a e ,  the kinoprene-  tr^a't'ed i n s e c t s a l s o e x h i b i t e d ||afval^.chaKac,feer.isteics.. ./  ;  O v e r 80% o f t h e t h i r d i n s t a r a l a t i f o r m nymphs t r e a t e d  with  k i n o p r e n e underwent a supernumerary moult b u t developed adults was  with  many a l a t e c h a r a c t e r i s t i c s .  The f o u r t h  the l e a s t s e n s i t i v e stage as v i r t u a l l y  into  instar  a l l the treated  71  insects  developed  When 65 ppm  into normal-looking alate virginoparae.  k i n o p r e n e was  applied  t o comparable  stages of  a p t e r i f o r m nymphs, 90% m o r t a l i t y  occurred within  treatment.  for this will  in  Although  a later  caused  no  section,  the reasons  i t s h o u l d be n o t e d  that  24 h r s . o f  be d i s c u s s e d  10  ppm  supernumerary m o u l t i n g o r d e v i a t i o n s  kinoprene  from  normal  development. These r e s u l t s c a n be  used  clearly  demonstrate  as JH m i m i c t o a l t e r  the epidermal c e l l s  i n alatiform  the normal nymphs.  o f a r t i f i c i a l l y e l e v a t e d JH t i t e r s , f o r m a c u t i c l e w h i c h was pearance. thesis  T h e r e f o r e , the  o f White  (1968b) and E l l i o t t  development o c c u r s i n the presence more, t h e d i f f e r e n t i a l instars the  to kinoprene  first  inhibiting  and  second  responses  the  final  1970;  nymphal  influence  were i n d u c e d  or apterous  i n ap-  (1975) t h a t o f low  JH  alatiform  levels.  Further-  alatiform  t h a t e l e v a t e d JH  levels  instars  are p a r t i c u l a r l y  important i n  during  f e a t u r e s and f a v o u r i n g  characteristics.  findings  Gilbert  instar  to  s u p p o r t the g e n e r a l hypo-  in  p e r s i c a e appear  opposed t o those o b t a i n e d i n o t h e r  (Wigglesworth  of  suggest  the r e t e n t i o n o f apterous  metrically  Under t h e  of the four  the development o f a l a t e  Additional  kinoprene  programming  the c e l l s  more j u v e n i l e findings  that  1975).  i s usually  dia-  exopterygotes  In the l a t t e r  species,  t h e most J H A - s e n s i t i v e  72  stage. the  However, i n  final  v i c i a e ( L e e s 1980) a n d  persicae,  f o u r t h n y m p h a l i n s t a r was a l m o s t t o t a l l y i n s e n s i t i v e  t o k i n o p r e n e a s t h e t r e a t e d nymphs d e v e l o p e d i n t o normal a l a t e a d u l t s . levels during  This  s u g g e s t s t h a t a d e c l i n e i n JH  t h e l a s t nymphal i n s t a r i s n o t p r e r e q u i s i t e t o  metamorphosis i n a l a t i f o r m aphids. the  I n an a t t e m p t t o e x p l a i n  a n o m a l o u s r e s p o n s e o f a p h i d s t o JHA, L e e s  that  apparently  (1) a d u l t d e t e r m i n a t i o n  i s virtually  (19 80)  proposed  c o m p l e t e by t h e  f i r s t d a y o f t h e l a s t i n s t a r a n d (2) more i m p o r t a n t l y , in  f o u r t h i n s t a r are i n s e n s i t i v e t o JH.  t h i s represented to regulate  an a d a p t a t i o n  i n time.  that the epidermal c e l l s  earlier  JH  separating  The f l a w i n t h e L e e s ' h y p o t h e s i s  is  i n the fourth i n s t a r f L persicae are  J H - s e n s i t i v e b e c a u s e when k i n o p r e n e was a p p l i e d t o  i n s t a r s , t h e c e l l s w e r e r e - p r o g r a m m e d t o p r o d u c e an  a d d i t i o n a l l a r v a l c u t i c l e and undergo a supernumerary before  that  i n aphids which permitted  b o t h metamorphosis and p o l y m o r p h i s m by  these functions  obviously  Lees claimed  tissues  p r o d u c i n g an a d u l t o i d c u t i c l e .  l i k e l y explanation  Therefore,  moult  a more  c a n be p r o p o s e d t o e x p l a i n t h e a p p a r e n t  i n s e n s i t i v i t y of t i s s u e s i n the fourth i n s t a r to kinoprene. In the reproduction  studies, kinoprene s i g n i f i c a n t l y  l a r v i p o s i t i o n rates of precocene-treated t h i s increase  virginoparae.  was n o t i m m e d i a t e b u t r a t h e r o c c u r r e d  d a y s a f t e r k i n o p r e n e was a p p l i e d .  This  increased  d e l a y may  However,  1 to 2 represent  73  the  time required  f o r kinoprene to d i f f u s e through the  and  reach s u f f i c i e n t l e v e l s to stimulate  In f o u r t h i n s t a r a l a t i f o r m s of M.  persicae,  t h i s delay  viciae  w o u l d be  stadium l a s t s 2 to 3 days.  critical  T h e r e f o r e , by  had  reached s u f f i c i e n t l e v e l s w i t h i n the  may  have been a l r e a d y The  Unlike  (Bowers e t a l . 1976;  persicae  d i d not  as t h e the  and  fourth  time  kinoprene  i n s e c t s , the adult  position rates declined f i r s t and  cells  cuticle.  precocene  studies.  Pener et a l . 1978),  to apteriform  nymphs o f  induce p r e c o c i o u s metamorphosis.  s u b s t a n t i a l i m p a i r m e n t o f CA  p r e c o c e n e as  tissues.  ( L e e s 1980)  second anomaly o c c u r r e d i n the  other species  target  c o m m i t t e d t o p r o d u c e an  precocene treatments administered M.  the  cuticle  However,  a c t i v i t y o c c u r r e d because  significantly  larvi-  i n insects treated  s e c o n d i n s t a r nymphs.  with  I t i s noteworthy  t h a t l a r v i p o s i t i o n c o n t i n u e d a l b e i t a t a much r e d u c e d  rate.  This  insects  suggests that  s u f f i c i e n t JH was  to permit l i m i t e d reproductive precocious metamorphosis. the  pea (c)  aphid  a c t i v i t y but  the  impair in  (MacKauer e t a l . 1 9 7 9 ) .  Sclerotization  i s known t o i n f l u e n c e  various  to  S i m i l a r f i n d i n g s were r e p o r t e d  In a d d i t i o n to r e g u l a t i n g JH  present i n these  s t u d i e s , JH  has  the  molting  and  a r c h i t e c t u r e of the  b e e n shown t o f a v o u r t h e  l a m e l l a t e - a p p e a r a n c e of the  metamorphosis, cuticle.  In  retention  of  l a r v a l endocuticle  and  influence  74  t y r o s i n e metabolism important  ( N e v i l l e 1975).  i n the current context  The l a t t e r e f f e c t i s  i n that the s c l e r o t i z i n g  a g e n t , N a c e t y l dopamine, and t h e major c u t i c u l a r melanin,  are derivatives of tyrosine.  In other  s y n t h e s i s i s r e g u l a t e d by ecdyson and b u r s i c o n In  persicae, extensive  pigment,  species,  ( N e v i l l e 1975) .  s c l e r o t i z a t i o n and m e l a n i z a t i o n  shortly after ecdysis i n alate adults.  If  s c l e r o t i z a t i o n and m e l a n i z a t i o n i n a p t e r o u s  occur  reduced a d u l t s c a n be  a t t r i b u t e d t o h i g h JH t i t e r s d u r i n g t h e nymphal s t a g e s , the processes  their  then  s h o u l d be s u p p r e s s e d when a l a t i f o r m nymphs a r e  t r e a t e d w i t h JHA.  T h i s was c o n f i r m e d  as kinoprene  to f i r s t ,  second, t h i r d ,  inhibited  s c l e r o t i z a t i o n and m e l a n i z a t i o n .  applications  a n d f o u r t h i n s t a r a l a t i f o r m nymphs  f i n d i n g s have been r e p o r t e d  i n other  aphid  Although species  similar (Mittler  e t a l . 1976; L e e s 1 9 7 7 , 1 9 8 0 ) , i t h a s n o t b e e n e s t a b l i s h e d whether t h e i n h i b i t o r y e f f e c t s o f JH a r e d i r e c t  (viz. arising  f r o m e f f e c t s on t h e c u t i c l e o r t y r o s i n e m e t a b o l i s m ) o r i n d i r e c t (viz.  i n h i b i t i o n o f bursicon or ecdyson). (d)  Cephalic sensory I n many a p h i d  and  sensory  sensilla)  antennal  are present  (Palmer 1952).  structures  s p e c i e s , i n c l u d i n g M^ p e r s i c a e ,  sensoria  (secondary  ocelli  rhinaria, placoid  i n a l a t e a d u l t s but absent i n apterae  Unlike previous  alate features, the effect of  JH on t h e d e v e l o p m e n t a n d d i f f e r e n t i a t i o n o f t h e s e  o  sensory  75  structures i s conflicting.  In  p e r s i c a e , t h e o c e l l i and  p a r t i c u l a r l y t h e a n t e n n a l p l a c o i d o r g a n s w e r e much easily  less  s u p p r e s s e d b y t o p i c a l JHA a p p l i c a t i o n s t h a n o t h e r  alate characteristics hydropene-  ( L e e s 1977, 1980).  In the l a t t e r  or kinoprene-treated insects which e x h i b i t e d a  maximal m o r p h o l o g i c a l response l a c k e d o c e l l i abnormal  study,  p l a c o i d organs.  and had s m a l l  A f t e r examining the e f f e c t s of the  JHA o n a l l p o s t n a t a l s t a g e s , L e e s  f o u n d no e v i d e n c e o f a  J H - s e n s i t i v e s t a g e and c o n c l u d e d t h a t b o t h s t r u c t u r e s were relatively the  i n s e n s i t i v e t o JH.  I n A ^ f a b a e a n d M^ p e r s i c a e ,  development o f t h e secondary a n t e n n a l s e n s i l l a and o c e l l i  w e r e n o t a f f e c t e d when f o u r t h i n s t a r p r e s u m p t i v e w e r e r e a r e d on p l a n t s s p r a y e d w i t h k i n o p r e n e 1976).  gynoparae  (Mittler et a l .  However, when t h e a l a t i f o r m o f f s p r i n g o f t h e s e  were a l s o m a i n t a i n e d on k i n o p r e n e - t r e a t e d f o l i a g e , developed i n t o a d u l t s which lacked o c e l l i secondary s e n s o r i a . of  females  they  a n d h a d f e w e r o r no  Therefore, contrary t o the conclusion  Lees, the f i n d i n g s o f M i t t l e r e t a l . suggest t h a t the  s t r u c t u r e s a r e s e n s i t i v e t o J H a t some p a r t i c u l a r  stage(s).  T h i s was c o n f i r m e d i n t h e p r e s e n t s t u d y a s t h e t i m i n g o f kinoprene a p p l i c a t i o n t o the various postnatal s i g n i f i c a n t e f f e c t on o c e l l a r development.  s t a g e s had a  A s r e p o r t e d by  M i t t l e r e t a l . (197 6), f o u r t h i n s t a r a l a t i f o r m  larvae,  were t r e a t e d w i t h k i n o p r e n e , d e v e l o p e d i n t o a d u l t s w i t h  which almost  76  normal o c e l l i .  When t r e a t e d t h i r d  adults, their o c e l l i  i n s t a r s developed into  were s m a l l e r and m a l f o r m e d .  In contrast,  when k i n o p r e n e was a p p l i e d t o f i r s t a n d s e c o n d i n s t a r  alati-  f o r m s , o c e l l a r d e v e l o p m e n t was a l m o s t c o m p l e t e l y s u p p r e s s e d . T h i s s u g g e s t s t h a t e l e v a t e d JH l e v e l s d u r i n g t h e s e s t a g e s (such as i n a p t e r i f o r m s ) would i n h i b i t t h e development o f the  ocelli.  As d e s c r i b e d i n  viciae  (Lees 1980), t h e  s e n s o r y a n t e n n a l s e n s o r i a were t h e l e a s t J H - l a b i l e c h a r a c t e r i s t i c examined i n t h i s  study.  alate  When k i n o p r e n e was  a p p l i e d t o a l l f o u r p o s t n a t a l stages, the development o f t h e s e n s o r i a was a b n o r m a l b u t n o t t o t a l l y  inhibited.  As abnor-  m a l i t i e s became p r o g r e s s i v e l y more p r o n o u n c e d when y o u n g e r i n s t a r s were t r e a t e d , t h e p r e s e n t f i n d i n g s s u g g e s t t h a t  per-  haps e l e v a t e d p r e n a t a l JH l e v e l s a r e i m p o r t a n t i n s u p p r e s s i o n of  the structures.  et  al.  (197 6)  T h i s w o u l d p e r h a p s e x p l a i n why  Mittler  were a b l e t o p r e v e n t t h e d e v e l o p m e n t o f t h e  s e n s o r i a when b o t h m o t h e r s a n d o f f s p r i n g w e r e r e a r e d o n kinoprene-treated (e)  Wing In  foliage,  development  D o r a l i s fabae  (Von Dehn 1 9 6 3 ) ,  brassicae  ( W h i t e 1968a; W h i t e a n d Lamb 1968) a n d t h e a n d r o c y c l i c of  persicae  (Mittler et al.  strain  1976), apterous v i r g i n o p a r a e  which would normally give b i r t h t o a l a t e o f f s p r i n g  were  i n d u c e d t o p r o d u c e a p t e r o u s o f f s p r i n g when t r e a t e d w i t h JHA.  77  The  c u r r e n t study s u b s t a n t i a t e s these  t h a t h i g h JH t i t e r s  i n the maternal  findings  and  haemolymph and  suggests early  nymphal s t a g e s i n h i b i t w i n g d e v e l o p m e n t and  favour  the  p r o d u c t i o n o f a p t e r o u s morphs.  When f o u r t h  instar  alatiform  nymphs were t r e a t e d w i t h 65 ppm  kinoprene,  into  a l a t e s w i t h n o r m a l - s i z e d wings.  applied  to t h i r d  wings.  The  i n s t a r produced  alatiforms.  A l t h o u g h w i n g d e v e l o p m e n t was  i n neonate  during postnatal  larvae,  a p t e r o u s a d u l t s w h i c h had  crowded u n t i l  75%  and  produced  not t o t a l l y  administered to  inhibit  a d u l t emergence.  of the o f f s p r i n g i n t o normal  Although  apterae.  by  wing  newly-ecdysed conditions.  t h r e e d a y s were  also  conditions resulted  the JHA-treated  over  adults  Even though c o n d i t i o n s  favoured a l a t e production, e l e v a t e d maternal sulted  parti-  wing  apterae i n the c o n t r o l groups,  produced  sup-  morphs.  been r e a r e d under crowded first  instar  rudimentary.  development,  could potentially  d u r i n g the  t h e p r o d u c t i o n o f 10%  developed  second  t h e p r e n a t a l e f f e c t s o f JH on  d e v e l o p m e n t , k i n o p r e n e was  offspring  pronounced  favour the p r o d u c t i o n o f apterous  To d e t e r m i n e  in  treatments balloon-shaped  t h e w i n g b u d s i n e m e r g e n t a d u l t s were  f o r m a t i o n and  The  adults with  administered to f i r s t  T h u s , e l e v a t e d JH t i t e r s cularly  Similar  developed  e f f e c t s o f k i n o p r e n e were e v e n more  when t h e compound was  pressed,  they  i n the p r o d u c t i o n of apterous  insects.  JH  levels I n M_^  re-  persicae,  78  crowding of  adults  and  first  i n s t a r nymphs f a v o u r s  alate  production  (Bonnemaison 1 9 5 1 ) .  As  kinoprene  production  during  my  findings  suggest  B a s e d on  the h i s t o l o g i c a l  these  c r o w d i n g s u p p r e s s e s JH appearance of arrived  at  the  CA,  similar  stages,  synthesis. White  contradict  persicae.  Using a r t i f i c i a l  demonstrated t h a t c h l o r i n a t e d isomeric  JH  m i x t u r e had  administered  to adults  authors observed only they concluded JHA  that  no  nymphs o f  impaired  i n t h i s study  Second, the  quite  the  less  dosages employed  JH  (dissolved  i s very o i l - s o l u b l e , i t could a b s o r b e d by  and  an  Since  high  of  the  aphids  the  the method  First,  CFA  i n m e t h a n o l ) was  dissolve  insects.  has  than  (Lees 19 8 0 ) .  i n e x p e r i m e n t s on  artificial  the  mortality,  bioassay  criticism.  Third,  JH  be  authors  i n t h i s study appear q u i t e  the  not  surface  develop-  morphologically-active  to  and  Parafilm  (CFA)  persicae.  However, t h e  t o x i c to a d u l t  synthetic  acid  d e v e l o p m e n t and  i s open t o  (5-2 5 u l JHA/ml. m e t h a n o l ) . larvae,  wing  a b n o r m a l e f f e c t s i n d u c e d by  r e c e n t l y b e e n shown t o be k i n o p r e n e and  on  a p p a r e n t a p t e r i z i n g e f f e c t when  and  any  JHA  d i e t s , these  farnesoic  were m e r e l y p a t h o l o g i c a l .  employed  (1975)  t h o s e o f Applebaum e t a l .  a l s o e x a m i n e d the. e f f e c t s o f  ment i n  Elliott  that  conclusions.  These f i n d i n g s (1975) who  (1968b) and  inhibited alate  diet. i n the  Finally,  As  high  neonate applied  synthetic  Parafilm  a l l the  experi-  79  m e n t s w e r e p e r f o r m e d on  artificial  diet.  Although  the  s t u d i e s w e r e d e s i g n e d so t h a t a s i g n i f i c a n t number o f adults developed, only alates.  As  7-15%  of the  some f a c t o r i n t h e the  not  They p o s t u l a t e  surprising.  of events, aphid duction  c o n t r o l s developed  d i e t a p p e a r s t o be  alate formation,  conclusions  o f Applebaum e t a l . a r e that " i n the  normal  to high  titre  and  a p p l i c a t i o n o f e x o g e n o u s JH  i n t e r n a l h o r m o n a l m i l i e u by  r e s u l t i n g the As  abnormalities in  persicae,  rudiments JH.  specific t o an  m a x i m a l l e v e l o f e n d o g e n o u s JH m i g h t s e r v e o n l y the  pro-  of endogenous  D i v e r s i o n t o winged a d u l t s would then r e q u i r e 'initiators',  course  development i s d i r e c t e d towards the  d o r m a n t due  into  limiting  o f a p t e r o u s a d u l t s , a s i t u a t i o n where w i n g  a r e p r e s e n t and  alate  already  to  disrupt  an o v e r a b u n d a n c e o f  JH,  recorded". JHA  h a v e b e e n shown t o  inhibit  w i n g d e v e l o p m e n t when a p p l i e d t o a l a t i f o r m nymphs a n d / o r alate-producing Despite  apterous adults of  t h i s , L e e s c o n c l u d e d t h a t JH  i n wing determination m e t h o d s e m p l o y e d by their  L e e s may Since  the  quently  revealed  involved  However, t h e  bioassay  results  crowded mothers o f apterae which are  as a p t e r a e .  and  viciae indistin-  t h i r d i n s t a r , Lees found i t necessary  t r e a t t h e n e o n a t e l a r v a e and  1980).  directly  have i n f l u e n c e d t h e  u s u a l l y p r o d u c e b o t h a l a t e s and until  i s not  i n this species.  interpretation.  guishable  v i c i a e ( L e e s 1977,  to  d i s c a r d t h o s e t h a t were subseTherefore,  any  of the  JHA-treated  80  i n s e c t s w h i c h e x h i b i t e d a p t e r o u s c h a r a c t e r i s t i c s would have been e l i m i n a t e d from t h e r e p o r t e d r e s u l t s .  I n a d d i t i o n , when  L e e s e x a m i n e d t h e e f f e c t s o f JH on a d u l t a p t e r a e , o n l y females which produced the r e s u l t s .  On  e x c l u s i v e l y a p t e r a e were r e p o r t e d i n  t h i s b a s i s , Lees concluded t h a t maternal  JH l e v e l s a r e u n i m p o r t a n t  i n wing  formation.  T h i s may  i n t r o d u c e d e x p e r i m e n t a l b i a s because e x c l u s i v e a l a t e a p t e r o u s f o r m a t i o n r a r e l y o c c u r s even under normal  C.  have and  conditions.  LETHAL EFFECTS JHA  h a v e b e e n shown t o h a v e c o n s i d e r a b l e p o t e n t i a l  i n the c o n t r o l of aphids The  those  (Tamaki 1973;  H r d y 1974;  p r e s e n t i n v e s t i g a t i o n s have d e m o n s t r a t e d  s t a g e s and morphs o f to kinoprene.  The  persicae differ  compound was  Staal  t h a t the v a r i o u s  in their  sensitivity  q u i t e t o x i c t o most  nymphs p a r t i c u l a r l y t h e y o u n g e r i n s t a r s . e f f e c t s were i m m e d i a t e l y a p p a r e n t ,  1975).  Although  the m a j o r i t y of  m o r t a l i t y o c c u r r e d d u r i n g t h e supernumerary and  alatiform  lethal the  adult stages.  A p t e r i f o r m nymphs w e r e e v e n more s e n s i t i v e t o k i n o p r e n e . When t h e y w e r e d i p p e d i n a 65 ppm 100%  kinoprene  m o r t a l i t y o c c u r r e d w i t h i n 24 h r s .  appears  t o be due  t o the presence  i n a p t e r o u s morphs.  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N a t u r e (Lond.) 2 6 5 : 5 4 8 - 5 5 0 . R i v n a y , E. 19 37. M o i s t u r e a s t h e f a c t o r a f f e c t i n g w i n g dev e l o p m e n t i n t h e c i t r u s a p h i d , I o x a p t e r a a u r a n t i i Boy. B u l l . E n t . Res. 2 8 : 1 7 3 - 1 7 9 . Schoonveld, H. 1979. P r e c o c e n e - i n d u c e d c o l l a p s e and r e a b s o r p t i o n o f c o r p o r a a l l a t a i n nymphs o f L o c u s t a m i g r a t o r i a . E x p e r i e n t i a . 3 5 :3 6 3- 3 6 4 . S h i n j i , G.O. 1918. A c o n t r i b u t i o n to the p h y s i o l o g y of development i n a p h i d s . B i o l . B u l l . 35:95-116.  wing  S h u l l , A.F. 1928. D u r a t i o n o f l i g h t and t h e w i n g s o f t h e a p h i d Macrosiphum s o l a n i f o l i i . A r c h . Entwicklungsmech. Organ. • 115:825-851. S h u l l , A.F. 1938. Time o f d e t e r m i n a t i o n and t i m e o f d i f f e r e n t i a t i o n of aphid wings. Amer. N a t . 7 2 : 1 7 0 - 1 7 9 .  85  S t a a l , G.B. 1 9 7 5 . I n s e c t growth r e g u l a t o r s w i t h j u v e n i l e hormone a c t i v i t y . Ann. Rev. E n t . 2 0 : 4 1 6 - 4 6 0 S u t h e r l a n d , O.R.W. 1 9 6 9 . The r o l e o f c r o w d i n g i n t h e p r o d u c t i o n o f a l a t e f o r m s b y two s t r a i n s o f t h e p e a a p h i d , A c y r t h o s i p h o n pisum. J . I n s e c t P h y s i o l . 1 5 : 1 3 8 5 - 1 4 1 0 . S u t h e r l a n d , O.R.W. 1 9 7 0 . An i n t r i n s i c f a c t o r i n f l u e n c i n g a l a t e p r o d u c t i o n by two s t r a i n s o f t h e p e a a p h i d , A c y r t h o s i p h o n pisum. J . I n s e c t P h y s i o l . 16:1349-1350. T a m a k i , G. 1 9 7 3 . Insect developmental i n h i b i t o r s : e f f e c t o f r e d u c t i o n and d e l a y c a u s e d b y j u v e n i l e hormone m i m i c s on t h e p r o d u c t i o n o f w i n g e d m i g r a n t s o f Myzus p e r s i c a e (Homoptera: A p h i d i d a e ) o n p e a c h t r e e s . Can. E n t . 1 0 5 : 761-765.  T o b a , H.H., J.D. P a s c h k e a n d S. F r i e d m a n 1 9 6 7 . Crowding as t h e p r i m a r y f a c t o r i n t h e p r o d u c t i o n o f t h e gamic a l a t e form o f T h e r i o a p h i s muculata (Homoptera: A p h i d i d a e ) . J. Insect P h y s i o l . 13:381-391. T u r n e r , W.F., and A.C. B a k e r 1 9 1 6 . On o c c u r r e n c e o f a n i n t e r mediate i n A p h i s pomi. E n t . S o c . Wash. P r o c . 17:4 2 - 5 2 . U n n i t h a n , G.C. and K.K. N a i r 1 9 7 9 . The i n f l u e n c e o f c o r p u s a l l a t u m a c t i v i t y o n t h e bug O n c o p e l t u s f a s c i a t u s t o p r e c o c e n e 1, 2, 3. Ann. E n t . S o c . Am. 7 2 : 3 8 - 4 0 . U n n i t h a n , G . C , K.K. N a i r , and W.S. Bowers 1 9 7 7 . Precoceneinduced d e g e n e r a t i o n o f the corpus a l l a t u m o f a d u l t f e m a l e s o f t h e bug O n c o p e l t u s f a s c i a t u s . J . I n s e c t P h y s i o l . 23:1081-1094. U n n i t h a n , G . C , K.K. N a i r , and A. S y e d 1 9 8 0 . Precocene-induced metamorphosis i n the d e s e r t l o c u s t S c h i s t o e e r c a g r e g a r i a . E x p e r i e n t i a . 36:135-136. Von  Dehn, M. 1 9 6 3 . Hemmung d e r G l u e g e l b i l d u n g d u r c h F a r n e s o l b e i der Schwarzen Bohnenlaus, D o r a l i s fabae Scop. N a t u r w i s s . 50 : 5 7 8 - 5 7 9 .  Wadley, F.M. 1 9 2 3 . F a c t o r s a f f e c t i n g the p r o p o r t i o n o f a l a t e and a p t e r o u s f o r m s o f a p h i d s . Ann. E n t . S o c . Am. 1 6 : 2 7 9 303.  Wellington, dusk.  W.G. 1 9 7 4 . Bumblebee o c e l l i and S c i e n c e (Wash.) 1 8 3 : 5 5 0 - 5 5 1 .  navigation a t  86  W h i t e , W.S. 1946. The e n v i r o n m e n t a l c o n d i t i o n s a f f e c t i n g t h e g e n e t i c mechanism o f w i n g p r o d u c t i o n i n t h e chrysanthemum aphid. Am. N a t . 80:245-270. W h i t e , D. 19 65. Changes i n s i z e o f t h e c o r p u s a l l a t u m p o l y m o r p h i c i n s e c t . N a t u r e (Lond.) 208:807.  in a  W h i t e , D. 1968a. P o s t n a t a l t r e a t m e n t o f t h e cabbage a p h i d w i t h a s y n t h e t i c j u v e n i l e hormone. J . Insect Physiol. 14:901-912. W h i t e , D. 19 6 8b. Cabbage a p h i d : E f f e c t o f i s o l a t i o n on f o r m and e n d o c r i n e a c t i v i t y . S c i e n c e (Wash.) 159:218-219. W h i t e , D., and K.P. Lamb 1968. Effects of a synthetic j u v e n i l e hormone on a d u l t c a b b a g e a p h i d s and t h e i r J . I n s e c t P h y s i o l . 14:395-402.  progeny.  W h i t e , D. 1971. Corpus a l l a t u m a c t i v i t y a s s o c i a t e d w i t h d e v e l o p m e n t o f w i n g b u d s i n c a b b a g e a p h i d embryos and larvae. J . I n s e c t P h y s i o l . 17:761-773. W i g g l e s w o r t h , V.B. Edinburgh.  1970.  I n s e c t hormones.  Oliver  and Boyd,  87 A p p e n d i x l a ( r e f e r s t o T a b l e 5) E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y nymphs o f a p t e r o u s a d u l t s  of f i r s t  instar  apteriform  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  3  4  5  4  4  3  3  2  31  3.1  2  0  4  3  3  3  4  3  4  3  3  30  3.0  3  0  3  4  4  5  5  3  4  4  2  35  3.5  4  2  3  4  3  4  4  3  3  3  4  33  3.3  5  0  3  3  4  4  3  5  3  4  2  31  3.1  6  0  2  4  3  5  3  4  4  3  3  31  3.1  Total Mean S.E.  3 0.5 0.3  18 3.0 0.2  21 3.5 0.2  21 3.5 0.2  26 4.3 0.3  23 3.8 0.3  22 3.C 0.3  21 3.5 0.2  20 3.3 0.2  16 2.6 0.3  191 31.8 0.78  19.1' 3.18 0.18  Total  Mean  DAYS # INSECT  Total  Mean  Kinoprene-- t r e a t e d 1  2  3  4  5  6  7  8  9  10  1  0  3  3  4  3  6  5  3  4  2  33  3.3  2  0  3  4  3  5  4  3  4  3  3  32  3.2  3  0  2  4  3  3  5  5  3  4  2  31  3.1  4  2  3  3  4  5  5  4'  3  3  3  36  3.6  5  1  3  4  2  6  3  4  3  2  4  32  3.2  6  1  2  4  3  5  3  4  4  2  2  31  3.1  Total 4 Mean 0.66 S.E. 0.33  17 22 19 27 26 25 20 18 16 195 2.66 3.66 3.16 4.50 4.33 4.16 3.33 3.00 2.66 32.5 0.21 0.21 0.30 0.50 0.49 0.33 0.32 0.39 0.16 0.76  19.5 3.25 0.07  Appendix  l b ( r e f e r s t o T a b l e 5)  E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y nymphs o f a p t e r o u s a d u l t s DAYS # INSECT  o f second  instar  apteriform  T o t a l . Mean  Control 1  2  3  4  5  6  7  8  9  10  1  0  3  4  4  5  5  4  3  3  2  33  3.3  2  0  4  3  4  4  6  3  4  4  2  34  3.4  3  0  3  4  3  4  *  14  2.8  4  0  3  3  4  4  4  5  3  3  3  32  3.2  5  2  2  4  3  3  5  4  4  4  3  34  3.4  6  0  3  4  5  5  4  4  3  2  3  33  3.3  18 3.00 0.21  22 3.66 0.21  23 3.80 0.30  25 4.16 0.30  24 4.80 0.37  20 4.00 0.31  17 3.40 0.24  Total 2 Mean 0 .33 S.E. 0 .33  16 13 3.20 2.60 0.37 0.24  180 30 3.21  19.38 3.23 0.09  Total  Mean  Kinoprene-- t r e a t e d  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  2  3  4  5  5  3  4  4  3  33  3.3  2  0  2  2  4  3  6  4  3  3  2  29  2.9  3  2  3  4  4  5  4  3  4  2  4  35  3.5  4  1  4  3  5  4  6  4  3  3  3  36  3.6  5  0  2  4  5  4  3  4  3  2  4  31  3.1  6  0  4  3  3  6  3  4  3  3  3  33  3.3  17 2.83 0.40  19 3.16 0.30  25 4.16 0.30  27 4.50 0.42  27 4.50 3.56  22 3.66 0.21  20 3.33 0.21  Total 3 Mean 0 .50 S.E. 0 .34 Note:  * denotes  death  17 > 19 197 2.83 3.16 32.83 0.30 0.30 1.04  19.7 3.28 0.10  89 Appendix ±  c  E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y nymphs o f a p t e r o u s a d u l t s  ( r e f e r s t o T a b l e 5)  of third instar  apteriform  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  4  4  5  6  4  3  4  3  36  3.6  2  0  4  3  4  6  5  4  4  3  3  36  3.6  3  0  2  4  3  4  5  3  3  3  2  29  2.9  4  0  3  3  5  4  6  4  3  3  3  34  3.4  5  2  3  *  5  2.5  6  2  4  3  4  3  6  4  3  2  4  35  3.5  19 3.16 0.30  17 3.40 0.24  20 4.00 0.31  22 4.40 0.50  28 5.60 0.24  19 3.80 0.20  16 3.20 0.20  15 3.00 0.31  15 3.00 0.31  4 Total Mean 0 .66 S.E. 0 .42  Total  Mean  175 19.50 29.16 3.25 4.94 0.18  Kinoprene-- t r e a t e d DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  4  5  4  4  3  4  2  4  33  3.3  2  0  3  3  3  6  3  4  2  3  3  30  3.0  3  1  2  4  4  5  3  4  3  3  3  31  3.1  4  0  3  3  5  4  5  3  4  4  2  33  3.3  5  2  2  4  3  5  4  3  3  2  3  31  3.1  6  0  3  3  4  4  5  2  4  2  2  29  2.9  16 2.66 0.21  21 3.50 0.22  24 4.00 0.36  28 4.66 0.33  24 4.00 0.36  19 3.16 0.30  20 3.33 0.33  16 2.66 0.33  Total 3 Mean 0.50 S.E. 0.37  Note: * denotes  death  Total  Mean  17 187 18.69 2.83 31.16 3.11 0.07 0.30 0.65  90 A p p e n d i x I d ( r e f e r s t o T a b l e 5) E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y nymphs o f a p t e r o u s a d u l t s  of fourth instar  apteriform  Total  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  2  4  3  5  4  3  2  2  28  2.8  2  0  2  4  3  5  3  6  4  3  2  32  3.2  3  0  3  2  4  4  6  3  4  2  4  32  3.2  4  0  2  4  3  5  4  5  3  4  3  33  3.3  5  2  2  3  4  4  3  6  4  2  4  34  3.4  6  2  3  4  4  5  4  3  2  5  3  34  3.4  Total Mean S.E.  4 0.66 0.42  22 3.66 0.21  26 4.33 0.33  20 3.33 0.33  18 3.00 0.51  18 3.00 0.51  193 32.16 0.90  19.3 3.21 0.09  15 19 2.50 3.16 0.22 0.40  25 27 4.16 4.50 0.47 0.56  Mean  Kinoprene-- t r e a t e d  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  4  3  5  4  3  4  2  3  31  3.1  2  0  4  3  4  3  6  4  3  2  4  33  3.3  3  0  3  4  3  3  5  4  3  2  3  30  3.0  4  0  2  4  3  3  6  3  2  4  3  30  3.0  5  2  2  4  4  5  3  4  4  3  2  33  3.3  6  1  4  3  3  6  4  3  2  4  4  33  3.3  18 3.00 0.37  22 3.60 0.21  20 3.30 0.21  25 4.16 0.54  18 3.00 0.36  17 2.80 0.46  19 3.10 0.30  190 31.6 0.61  19 3.16 0.06  Total Mean S.E.  3 0.50 0.34  28 21 4.60 3.50 0.19 . 0.24  Total  Mean  91 Appendix  2 ( r e f e r s t o F i g . 15)  E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y o f n e w l y - e c y s e d adults  apterous  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  2  3  3  4  5  5  3  4  3  2  34  3.4  2  0  4  3  4  4  5  3  4  2  3  32  3.2  3  2  3  4  2  6  4  3  3  4 •  2  33  3.3  4  0  4  3  4  3  6  4  3  3  3  33  3.3  5  0  3  4  4  5  5  4  4  3  2  35  3.5  Total Mean S.E.  4 0.80 0.48  17 3.40 0.24  17 3.40 0.24  18 3.60 0.40  23 4.60 0.50  25 5.00 0.31  17 3.40 0.24  17 3.60 0.24  15 3.00 0.31  12 3.40 0.24  167 33.4 0.50  16.7 3.34 0.05  Total  Mean  Total  Mean  Kinoprene-- t r e a t e d  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  2  4  4  5  5  3  4  4  2  33  3.3  2  0  3  4  5  4  6  2  4  3  3  34  3.4  3  0  2  3  4  4  6  3  4  2  4  32  3.2  4  2  3  4  2  5  3  4  2  2  4  31  3.1  5  1  4  4  4  2  6  3  4  3  3  34  3.4  Total Mean S.E.  3 0.60 0.40  14 2.80 0.37  19 3.80 0.20  19 3.80 0.48  20 4.00 0.54  18 3.60 0.40  14 2.80 0.37  16 164 3.20 32.8 0.37 0.58  16.4 3.28 0.05  26 15 5.20 3.00 0.58 0.31  92 Appendix  3a ( r e f e r s t o T a b l e 6)  E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y nymphs o f a l a t e a d u l t s  of first  instar  apteriform  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  2  3  3  4  3  3  2  3  2  25  2.5  2  0  3  3  4  4  5  2  3  2  3  29  2.9  3  1  2  3  2  5  3  3  4  2  4  29  2.9  4  1  3  3  4  4  4  3  4  3  4  33  3.3  5  2  2  3  4  3  5  4  2  3  2  31  3.1  Total Mean S.E.  4 0.80 0.35  12 2.40 0.24  15 3.00 0.00  17 3.40 0.40  20 4.00 0.31  20 4.00 0.44  15 3.00 0.31  15 3.00 0.44  13 2.60 0.24  15 3.00 0.44  147 29.4 1.32  14.7 2.94 0.13  Total  Mean  DAYS # INSECT  Total  Mean  Kinoprene-- t r e a t e d 1  2  3  4  5  6  7  8  9  10  1  0  2  3  4  3  3  4  2  3  2  26  2.6  2  0  3  3  4  4  2  4  3  3  2  28  2.8  3  1  2  3  4  3  3  4  2  3  3  28  2.8  4  0  4  3  5  3  4  3  4  4  2  32  3.2  5  1  2  4  3  5  3  4  3  3  2  30  3.0  Total Mean S.E.  2 0.40 0.24  13 2.60 0.46  16 3.20 0.20  20 4.00 0.31  18 3.60 0.40  15 3.00 0.31  19 3.80 0.20  14 2.80 0.37  16 3.20 0.20  11 3.20 0.20  144 28.8 1.01  14.4 2.88 0.10  93 Appendix  3b ( r e f e r s t o T a b l e 6)  E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y o f s e c o n d nymphs o f a l a t e a d u l t s  instar  apteriform  Control DAYS  Total  Mean  1  2  3  4  5  6  7  8  9  10  1  0  1  3  4  4  5  4  3  2  3  29  2.9  2  2  3  3  5  6  4  4  3  2  2  34  3.4  3  0  2  4  3  4  6  3  4  3  3  32  3.2  4  2  1  5  3  4  5  3  3  2  4  32  3.2  5  1  4  3  5  6  3  4  4  3  3  36  3.6  23 4.60 0.50  18 3.60 0.15  17 3.40 0.24  12 2.40 0.24  15 3.00 0.31  163 32.6 1.16  16.3 3.26 0.11  Total  Mean  #  INSECT  Total Mean S.E.  5 1.00 0.44  11 2.20 0.58  18 3.60 0.40  20 4.00 0.44  24 4.80 0.48  Kinoprene-- t r e a t e d DAYS 1  2  3  4  5  6  7  8  9  10  1  0  2  3  4  4  3  3  2  4  2  27  2.7  2  2  3 •  4  3  5  3  4  3  3  3  33  3.3  3  0  3  4  3  5  4  3  2  3  2  29  2.9  4  0  4  2  5  4  3  4  4  2  4  32  3.2  5  0  2  4  3  4  5  3  4  4  2  31  3.1  Total Mean S.E.  2 0.40 0.40  17 3.40 0.40  18 3.60 0.40  13 2.60 0.40  152 30.4 1.07  INSECT  14 2.80 0.37  22 4.40 0.24  18 3.60 0.40  17 3.40 0.24  15 3.00 0.44  16 3.20 0.37  15.2 3.04 0.10  94 Appendix  3c ( r e f e r s t o T a b l e 6)  E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y o f t h i r d nymphs o f a l a t e a d u l t s  instar  apteriform  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  2  3  3  4  4  3  2  4  1  26  2.6  2  0  3  4  4  5  3  4  3  4  3  33  3.3  3  0  2  4  3  2  5  4  3  3  2  28  2.8  4  0  3  3  4  4  3  3  2  2  4  28  2.8  5  1  2  1  5  4  3  4  3  3  3  29  2.9  Total Mean S.E.  1 0.20 0.20  12 2.40 .0.24  15 3.00 0.54  19 3.80 0.37  19 3.80 0.48  18 3.60 0.40  13 2.60 0.50  144 28.8 1.15  14.4 2.88 0.11  Total  Mean  13 13 16 3.60 '' 2.60 3.20 0.24 0.24 0.37  Total  Mean  K i n o p r e n e -- t r e a t e d DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  '1  0  2  3  4  4  3  4  3  2  4  29  2.9  2  1  2  3  4  3  5  3  3  4  2  30  3.0  3  2  3  4  3  4  3  4  2  3  3  31  3.1  4  1  2  3  4  5  3  4  2  3  4  31  3.1  5  0  4  3  2  6  3  4  3  2  3  30  3.0  Total Mean S.E.  4 0.80 0.37  13 2.60 0.40  16 3.20 0.20  17 3.40 0.40  22 4.40 0.50  17 3.40 0.40  19 3.80 0.20  13 2.60 0.24  14 2.80 0.37  16 3.20 0.37  151 30.2 0.37  15.1 3.02 0.03  95 A p p e n d i x 3d ( r e f e r s t o T a b l e 6) E f f e c t s o f k i n o p r e n e on t h e f e c u n d i t y nymphs o f a l a t e a d u l t s  of fourth instar  apteriform  Control DAYS  1  2  3  4  5  6  7  8  9  10  1  0  3  3  4  3  4  2  3  3  2  27  2.7  2  0  3  3  2  5  3  4  3  2  3  28  2.8  3  1  2  3  4  5  3  4  2  4  2  30  3.0  4  1  3  4  4  3  5  2  3  4  4  33  3.3  5  0  3  2  4  3  5  4  3  3  3  30  3.0  Total Mean S.E.  2 0.40 0.24  14 2.80 0.20  15 3.00 0.31  18 3.60 0.40  19 3.80 0.48  20 4.00 0.44  16 3.20 0.48  14 2.80 0.20  16 3.20 0.37  14 2.80 0.37  . 148 29.6 1.02  14.8 2.96 0.10  #  INSECT  Total  Mean  Kinoprene-treated DAYS # INSECT  1  2  1  0  2  2  0 1  4  2  Total Mean S.W.  0  2  3 3 0.60 0.40  5  5 5 21 4.20 0.37  7 4  5 4  3  13 15 2.60 3.00 0'.24 0.31  5  4 3  2  6  4  4 3  3  4  3  3  3  5  3  3  3  4  3  20 4.00 0.54  4 3  4  3 3  2 17 3.40 0.24  9  2  4  2 4  8  10  Total  Mean  4  31  3.1  3  32  3.2  2  28  2.8  32  3.2  30  3.0  2 4  3 4  2  4  4  16 3.20 0.37  17 3.40 0.37  4 3  15 3.00 0.44  16 3.20 0.37  153 30.6 0.70  15.3 3.06 0.07  A p p e n d i x 4a, ( r e f e r s t o T a b l e 8) E f f e c t s of precocene-II apterous adults  on t h e  fecundity  of  first  instar  apteriform  nymphs  of  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  3  4  5  4  4  3  3  2  31  3.1  2  0  4  3  3  3  4  3  4  3  3  30  3.0  3  1  3  4  4  5  5  3  4  4  2  35  3.5  4  2  3  4  3  4  4  3  3  3  4  33  3.3  5  0  3  3  4  4  3  5  3  4  2  31  3.1  6  0  2  4  3  5  3  4  4  3  3  31  3.1  Total Mean S.E.  3 0.60 0.50  18 3.00 3.00  21 3.50 3.50  21 3.50 3.50  23 3.83 3.80  22 3.66 3.60  21 3.50 3.50  20 3.33 3.30  16 2.66 2.60  191 31.83 0.74  19.1 3.18 0.08  Total  Mean  26 4 . 33 4.30  Total  Mean  Precocene-treated DAYS # INSECT  1  1  2  3  4  5  6  7  8  9  10  0  1  1  0  2  0  1  1  1  1  8  0.8  2  0  1  0  1  2  1  0  1  0  6  0.6  3  0  1  1  2  0  2  1  0  1  0  7  0.7  4  0  0  2  1  1  1  1  0  0  2  8  0.8  5  0  2  1  0  1  1  0  0  1  1  7  0.7  6  0  1  2  2  0  0  2  1  1  0  9  0.9  Total Mean S.E.  0 0 0  7 1.16 0.30  6 1.00 0.36  6 1.00 0.36  .  6 1.00 0.25  Kinoprene  application  to  5 0.83 0.30  5 0.83 0.30  four-day-old  3 0.50 0.22  4 0.80 0.20  4 0.80 0.37  precocene-treated  45 7.50 0.42  4.5 0.75 0 . 04  insects  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  1  1  0  1  2  1  2  3  1  12  1.2  2  0  2  0  2  0  0  2  2  2  3  13  1.3  3  0  0  2  1  1  1  3  1  3  2  14  1.4  4  0  0  ' 1  1  2  3 .  1  3  2  1  14  1.4  5  0  2  1  0  1  2  2  3  1  3  14  1.4  6  0  0  2  1  *  3  0.7  Total Mean S.E.  0 0 0 Note:  5 0.83 0.40  7 1.16 0.30  * denotes  5 0.83 0.31 death  5 1.00 0.50  8 1.60 0.50  9 1.80 0.37  11 2.20 0.37  11 2.20 0.37  10 2 .20 0.44  Total  70 11.60 1.76  Mean  7.4 1.24 0.10  A p p e n d i x 4b ( r e f e r s t o T a b l e 8) E f f e c t s of precocene-II of apterous adults  on  the  fecundity  of  second i n s t a r  apteriform  nymphs  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  4  4  5  5  4  3  3  2  33  3.3  2  0  4  3  4  4  6  3  4  4  2  34  3.4  3  0  3  4  3  4  *  14  2.8  4  0  3  3  4  4  4  5  3  3  3  32  3.2  5  0  2  4  3  5  5  4  4  4  3 '  34  3.4  6  2  3  4  5  5  4  4  3  2  3  35  3.5  18 3.00 0.25  22 3.66 0.21  23 3.83 0.24  27 4.50 0.22  24 4.80 0.37  20 4.00 0.31  17 3.40 0.24  16 3.20 0.24  13 2.60 0.24  182 33.3 3.29  19.6 3.26 0.11  Total  Mean  Total Mean S.E.  2 0 . 33 0.33  Total  Mean  Precocene-treated DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  1  0  2  1  1  0  1  2  1  9  0.9  2  0  0  2  0  2  1  1  2  0  2  10  1.0  3  0  1  1  1  0  0  1  *  4  0.5  4  0  1  2  2  1  1  0  1  0  0  8  0.8  5  0  0  1  0  0  2  1  1  1  0  6  0.6  6  0  2  0  0  2  1  1  0  0  0  6  0.6  Total Mean S.E.  0 0 0  6 1.00 0.36  5 0.83 0.40  6 1.00 0.36  3 0.60 0.40  3 0.50 0.40  5 0.83 0 . 33  Kinoprene DAYS # INSECT  application  to  6 1.00 0.25  4 0.60 0.21  four-day-old  5 1.00 0 . 31  precocene-treated  43 7.16 0.90  4.4 0.73 0.08  insects  1  2  3  4  5  6  7  8  9  10  1  0  2  1  0  2  2  3  2  2  3  17  1.7  2  0  0  1  0  1  3  1  2  4  2  14  1.4  3  0  0  1  2  0  2  2  2  2  2  14  1.4  4  0  0  2  1  0  1  3  1  3  3  14  1.4  0  0  0  1  0  2  2  2  2  2  11  1.1  6  0  2  1  2  1  1  1  3  3  2  16  1.6  Total Mean S.E.  0 0 0  4 0.66 0.42  12 2.00 0.36  12 2.00 0.25  16 2.66 0.33  14 86 2 . 33 1 4 . 3 3 0.21 0.84  8.6 1.43 0.08  5  '  Note:  6 1.00 0.25  * denotes  6 1.00 0 . 36 death  4 0.66 0.33  11 1.83 0 . 30  Total  Mean  Appendix. 4 c ( r e f e r s t o Table 8 ) E f f e c t s of precocene-II of apterous adults  on the  fecundity  of  third  instar  apteriform  nymphs  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  4  4  5  6  4  3  4  3  36  3.6  2  0  4  3  4  6  5  4  4  3  3  36  3.6  3  0  2  4  3  4  5  3  3  3  2  29  2.9  4  0  3  3  5  4  6  4  3  3  3  34  3.4  5  2.5  35  3.5  175 29.16 4.94  19.5 3.25 0.18  5  2  3  *  6  2  4  3  Total Mean S.E.  4 0.60 0.42  19 3.16 0.30  17 3.40 0 . 31  4  3  6  4  3  20 4.00 0.50  22 4.40 0.24  28 5.60 0.20  19 3.80 0.20  16 3.20 0.20  2 15 3.00 0 . 31  4 15 3.00 0.31  Total  Mean  Precocene- •treated DAYS 1  2  3  4  5  6  7  8  9  10  1  0  2  3  2  1  0  0  3  2  2  15  1.5  2  0  0  1  2  2  3  0  2  3  2  15  1.5  3  0  0  1  1  3  2  3  1  1  0  12  1.2  4  0  *  5  0  4  3  2  2  2  3  2  2  3  23  2.3  6  0  3  4  3  3  2  3  2  3  0  23  2.3  Total Mean S.E.  0 0 0  9 1.80 0.80  12 2.40 0.60  10 2.00 0.31  11 2.20 0.37  9 1.80 0.48  9 1.80 0.73  10 2.00 0.31  11 2.20 0.37  7 1.40 0.60  INSECT  Kinoprene  Mean  0  application  DAYS  to  four - d a y - o l d  p r e c o c e n e! - t r e a t e d  1  2  3  4  5  6  7  8  9  10  1  0  2  1  2  1  3  2  4  4  2  0  0  3  1  2  4  3  3  3  0  3  1  3  4  3  3  4  0  1  2  2  4  4  3  INSECT  insect  3  22  2.2  3  2  21  2.1  4  3  3  27  2.7  3  2  3  24  2.4  5  1.6  25  2.5  124 20.66 3^.25  13.5 2.25 0.91  2  2  1  6  1  0  2  3  3  4  4  3  3  2  Total Mean S.E.  3 0.50 0.37  8 1.30 0.53  10 1.60 0.33  11 2.20 0.37  14 2.80 0.58  18 3.60 0.24  15 3.00 0.31  17 3.40 0.24  15 3.00 0.31  13 2.60 0.24  death  8.8 1.46 0 . 34  Mean  5  * denotes  88 14.66 3.47  Total  *  Note:  Total  99 A p p e n d i x 4d ( r e f e r s t o T a b l e 8) E f f e c t s of precocene-II of apterous adults  on t h e  fecundity  of  fourth  instar  a p t e r i f o r m nymphs  Control DAYS # INSECT  Total  Mean  1  2  3  4  5  6  7  8  9  10  1  0  3  2  4  3  5  4  3  2  2  28  2.8  2  0  2  4  3  5  3  6  4  3  2  32  3.2  3  0  3  2  4  4  6  3  4  2  4  32  3.2  4  0  2  4  3  5  4  5  3  4  3  33  3.3  5  2  2  3  4  4  3  6  4  2  4  34  3-4  6  2  3  4  4  5  4  3  2  5  3  35  3.5  Total Mean S.E.  4 0.60 0.46  15 2.50 0.34  19 3.10 0.43  22 3.60 0.23  25 4.10 0.38  25 4.10 0.52  27 4.50 0.61  20 3.30 0 . 36  18 3.00 0.56  194 18 3.00 32.33 0.98 0 . 39  19.4 3.23 0.09  Precocene- treated DAYS # INSECT  Total  Mean  1  2  3  4  5  6  7  8  9  10  1  0  3  2  3  3  4  2  4  3  3  27  2.7  2  1  3  3  2  3  3  4  3  4  1  27  2.7  3  2  2  1  4  2  4  2  4  2  3  26  2.6  4  0  1  2  2  3  3  4  3  2  2  22  2.2  5  0  2  4  1  4  3  4  2  4  2  26  2.6  6  0  2  2  3  3  3  3  3  2  3  24  2.4  Total Mean S.E.  3 0.50 0.43  13 2.16 0.33  14 2.30 0.46  15 2.50 0.46  18 3.00 0.28  20 3.30 0.21  19 3.10 0.43  17 2.80 0.38  16 2.60 0.45  14 2.30 0.36  152 25.33 0.80  15.2 2.53 0.08  Kinoprene  application  f o u r - d a y -•old p r e c o c e n e - t r e a t e d  to  DAYS  insects Total  Mean  1  2  3  4  5  6  7  8  9  10  1  0  2  3  2  3  5  4  3  2  3  27  2.7  2  0  1  2  3  4  3  5  3  3  2  26  2.6  3-  0  3  1  4  2  5  2  4  2  4  27  2.7  4  0  1  3  2  4  3  4  4  2  3  26  2.6  5  0  4  2  3  2  4  5  3  4  2  29  2.9  6  0  3  2  1  4  3  5  4  3  3  28  2.8  14 2.30 0.53  13 2.10 0.33  15 2.50 0.46  19 3.10 0.43  23 3.80 0.43  25 4.10 0.52  21 3.50 0.24  16 2.60 0.40  163 17 2.80 27.16 0.47 0 . 33  16.3 2.71 0.04  INSECT  Total Mean S.E.  0 0 • 0  100  A p p e n d i x 4e ( r e f e r s t o T a b l e 8) E f f e c t s o f p r e c o c e n e - I I on t h e apterous adults  fecundity  of  newly-- e c d y s e d  Control  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  2  3  3  4  5  5  3  4  3  2  34  3.4  2  0  4  3  4  4  5  3  4  2  3  32  3.2  3  2  3  4  2  6  4  3  3  4  2  33  3.3  4  0  4  3  4  3  6  4  3  3  3  33  3.3  5  0  3  4  4  5  5  4  4  3  2  34  3.4  6  0  4  2  4  3  3  4  4  3  2  "39  2.9  22 3.60 0.33  26 4.33 0.49  28 4.66 0.42  21 3.50 0.22  22 3.70 0.21  18 3.00 0.25  195 14 2.30 32.50 0.21 0.76  19.5 3.25 0.07  Total Mean S.E.  DAYS  4 21 19 0.66 3.50 3.16 0.42 0.22 0.30  Total  Mean  Precocene - t r e a t e d Total  Mean  1  2  3  4  5  6  7  8  9  10  1  0  0  1  2  1  1  0  2  0  0  7  0.7  2  0  2  0  1  0  2  1  1  0  1  8  0.8  3  0  0  2  0  2  1  1  0  1  0  .7  0.7  4  0  0  3  0  2  1  2  1  1  1  11  1.1  5  0  0  2  2  0  0  2  0  1  2  9  0.9  6  0  1  1  1  2  0  1  1  0  0  7  0.7  Total Mean S.E.  0 0 0  3 0.60 0.36  6 1.00 0.36  7 1.16 0.40  5 0.83 0.30  49 8.16 0.65  4.9 0.81 0.06  Insect  9 1.50 0.42  7 1.16 0.30  5 0.83 0.30  4 3 0.53 0.66 0.22 0.33  101  Appendix 4e-- continued  DAYS  K i n o p r e n e a p p l i c a t i o n t o p r e c o c e n e - t r e a t e d 24 h r s . o l d insects 1  2  3  4  5  6  7  8  9  10  1  0  0  2  0  2  0  3  4  2  3  16  1.6  2  0  0  2  1  2  1  4  2  3  2  17  1.7  3  0  1  1  3  0  3  2  3.  2  2  17  1.7  4  0  1  2  1  2  2  2  3  1  4  18  1.8  5  0  0  0  2  1  3  2  3  4  1  16  1.6  6  0  *  Total Mean S.E.  0 0 0  2 0.40 0.24  #  Total  Mean  INSECT  0 7 1.40 0.40  7 1.40 0.50  7 1.40 0.40  9 13 1.80 2.60 0.58 0.40  15 3.00 0.31  12 2.40 0.55  12 2.40 0.50  84 16.8 0.37  Kinoprene a p p l i c a t i o n t o precocene-treated 4-day-old DAYS  8.4 1.68 0.03  insects  Total  Mean  1  2  3  4  5  6  7  8  9  10  1  0  2  1  3  4  3  4  2  3  1  23  2.3  2  1  0  2  3  3  5  3  4  2  3  26  2.6  3  0  1  0  3  3  4  3  4  3  3  25  2.5  4  1  0  2  2  4  3  4  3  4  1  24  2.4  5  0  2  1  3  3  4  3  4  2  4  26  2.6  6  0  *  Total Mean S.E.  2 0.33 0.21  #  INSECT  0  14 5 6 1.20 1.00 2.80 0.44 0.37 0.20  Note: * denotes  17 3.40 0.24  death  19 17 3.80 3.40 0.37 0.24  17 3.40 0.40  12 14 2.80 2.40 0.37 0.60  124 24.8 0.58  12.4 2.48 0.05  102 A p p e n d i x 5a. ( r e f e r s t o T a b l e 9) E f f e c t s of precocene-II of a l a t e a d u l t s  on  the  fecundity  of  First  instar  apteriform  nymphs  Total  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1 '  0  2  3  3  4  4  3  3  2  1  25  2.5  2  2  1  4  3  3  4  2  3  3  2  27  2.7  3  0  2  4  3  4  3  4  3  4  1  28  2.8  4  0  3  2  4  4  3  2  3  3  2  26  2.6  5  2  2  1  3  4  2  3  4  2  4  27  2.7  6  0  0  1  *  1  0.3  10 2.00 0 . 54  134 2.23 4 .28  13.6 2 . 27 0.39  Total  Mean  Total Mean S.E.  4 0.66 0 . 42  10 1.60 0.42  15 2.50 0.56  16 3.20 0.20  19 3.80 0.20  16 3.20 0.37  14 2 .80 0.37  16 3.20 0.20  14 2.80 0.37  Mean  P r e c o c e n e - •treated 1 DAYS 1  2  3  4  5  6  7  8  9  10  1  0  1  2  2  0  0  1  0  2  0  8'  0.8  2  0  1  1  1  0  1  2  2  1  0  9  0.9  3  1  2  2  0  0  0  1  0  2  1  9  0.9  4  0  0  0  2  1  1  0  1  0  2  7  0.7  5  0  1  2  1  2  3  0  0  0  1  10  1.0  6  1  1  *  2  0.2  45 7.50 1.17  4.5 0.75 0.04  INSECT  Total Mean S.E.  2 0.33 0.21  6 1.00 0.25  Kinoprene  7 1.40 0.40  6 1.20 0 . 37  application  3 0.60 0.40  to  5 1.00 0.54  4 0.80 0.37  four-day-old  3 0.60 0.40  5 1.00 0.44  4 0.80 0 . 37  precocene-treated  insects  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  1  2  0  2  1  2  1  2  1  12  1.2  2  0  2  1  1  0  2  1  2  1  2  12  1.2  3  0  0  2  1  1  1  3  0  2  0  10  1.0  4  0  1  1  2  0  2  1  2  1  ' 1  11  1.1  5  0  1  0  1  2  1  1  2  0  2  10  1.0  6  0  *  Total Mean S.E.  0 0 0  5 1.00 0.31  Note:  6 1.20 0.37  * denotes  5 1.00 0.31 death  5 1.00 0.44  7 1.40 0.24  8 1.60 0.40  7 1.40 0.40  6 1.20 0.37  6 1.20 0.37  Total  55 11.00 0.44  Mean  5.5 1.10 0.04  103 A p p e n d i x .5b ( r e f e r s t o T a b l e 9) E f f e c t s of precocene-II of a l a t e adults  on  the  fecundity  of  second  instar  apteriform  nymphs  Control DAYS # INSECT  2  3  4  5  6  7  8  9  10  1  0  2  3  3  4  3  3  2  3  2  25  2.5  2  0  3  3  4  4  5  2  3  3  3  30  3.3  3  1  2  3  2  5  3  3  4  2  4  29  2.9  4  1  3  3  4  4  4  3  4  2  4  32  3.2  5  2  2  3  4  3  5  4  2  3  . 2  30  3.0  12 2.40 0.24  15 3.00 0.00  17 3.84 0.40  20 4.00 0.31  146 29.20 1.15  14.9 2.98 0.13  Total Mean S.E.  4 0 . 80 0.37  20 4 .00 0.44  15 3.00 0 . 31  15 3.00 0 .44  13 2.60 0.24  Total  Mean  1  15 3.00 0.44  P r e c o c e n e - •treated DAYS 9  10  2  0  1  0  8  0.8  1  0  1  0  1  7  0.7  1  0  1  2  0  1  9  0.9  0  2  1  2  0  1  1  8  0.8  2  1  1  1  1  0  0  10  1.0  2 0.40 0.24  3 0.60 0.24  42 8.40 0.50  4.2 0.84 0.05  2  3  4  5  6  7  1  0  2  1  0  0  2  2  0  1  0  1  2  3  0  1  2  1  4  0  0  1  5  1  2  1  '  Total  Mean  8  1 INSECT  Total Mean S.E.  2 0.20 0.20  6 1.20 0 . 37  Kinoprene  •  5 1.00 0 . 31  4 0.80 0 . 37  application  6 1.20 0 . 37  to  5 1.00 0.31  .  6 1.20 0.37  f o u r - d a y -•old  4 0 . 80 0 . 37  p r e c o c e n ei - t r e a t e d  insects  DAYS Total  Mean  1  2  3  4  5  6  7  8  9  10  1  0  1  2  0  3  1  2  3  0  3  15  1.5  2  0  1  1  2  0  3  0  2  1  2  12  1.2  3  0  0  2  0  3  0  3  1  2  3  14  1.4  4  1  1  0  1  1  2  3  1  2  0  12  1.2  5  0  1  1  2  0  2  2  2  3  1  14  1.4  Total Mean S.E.  1 0.20 0.20  8 1.60 0.50  10 2.00 0.54  INSECT  4 0.80 0.20  6 1.20 0.37  5 1.00 0.44  7 1.40 0.67  9 1.80 0 . 37  8 1.60 0.50  8 1.60 0.50  67 13 . 4 0 0.60  6.7 1..34 0.06  104 A p p e n d i x 5c ( r e f e r s t o T a b l e 9) E f f e c t s of precocene-II of alate adults  on t h e  fecundity  of  third  instar  apteriform  nymphs  Total  Control DAYS # INSECT  1  2  3  4  5  6  1  0  2  3  3  4  4  2  0  3  4  4  5  3  0  2  4  3  4  0  3  3  5  1  2  Total Mean S.E.  1 0.20 0.20  12 2.40 0.24  7  8  9  10  3  2  4  1  25  2.5  3  4  2  4  3  32  3.2  2  5  4  3  3  2  28  2.8  4  4  3  3  2  2  4  28  2.8  1  5  4  3  4  3  3  3  29  2.9  15 3.00 0.54  19 3.80 0.37  19 3.40 0.60  18 3.60 0.40  18 3.60 0.24  12 2.40 0.24  16 3.20 0.37  13 2.60 0.50  142 28.40 1.12  14.2 2.84 0.11  •  Mean  Precocene-treated DAYS 10  INSECT  Total  Mean  1  0  0  1  2  0  1  9  0.9  2  0  3  1  2  3  2  14  1.4  3  2  3  4  3  2  1  22  2.2  4  1  1  2  2  1  0  15  1.5  5  0  3  2  1  3  1  15  1.5  75 15 2.07  7.5 1.50 0.20  Total Mean S.E.  3 0 . 60 0.44  1.00 0.44  Kinoprene  1.80 0.37  10 2.00 0.63  application  10 2.00 0.89  to  10 2.00 0.31  9 1.40 0.24  -doy-old  1.80 0.58  1.40 0.50  precocene  5 1.00 0.37  treated  insects  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  2  1  1  2  3  4  4  ' 3  3  23  2.3  2  0  0  3  3  3  4  4  3  3  2  25  2.5  3  0  2  2  3  4  3  4  3  2  4  27  2.7  4  2  1  2  2  5  4  3  3  • 3  2  27  2.7  5  1  0  3  2  3  5  4  3  2  4  27  2.7  Total Mean S.E.  3 0.60 0.40  5 1.00 0.40  11 2.20 0.37  11 2.20 0.37  17 3.40 0.50  19 3.80 0.37  19 3.80 0.20  16 3.20 0.20  13 2.60 0.24  15 3.00 0.40  129 25.80 0.80  12.9 2.58 0.08  Total  Mean  105 A p p e n d i x 5d ( r e f e r s t o T a b l e 9) E f f e c t s of p r e c o c e n e - I I of a l a t e adults  on  the  fecundity  of  fourth  instar  apteriform  nymphs  Control DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  3  3  4  3  4  2  3  3  2  _27  2.7  2  0  3  3  2  5  3  4  3  2  3  28  2.8  3  1  2  3  4  5  3  4  2  4  2  30  3.0  4  1  3  4  4  3  5  2  3  4  4  33  3.3  5  0  3  2  4  3  5  4  3  3  3  30  3.0  Total Mean S.E.  2 0.40 0.24  14 2.80 0.44  18 3.60 0.40  19 3.80 0.48  20 4.20 0.48  14 2.80 0.20  16 3.20 0.37  14 2.80 0 . 37  148 29.6 1.02  14.8 2.96 0.10  Total  Mean  15 3.00 0 . 31  16 3 .20 0.48  Total  Mean  Precocene- •treated DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  0  2  3  2  2  3  3  2  2  19  1.9  2  0  2  0  1  3  4  2  4  2  4  22  2.2  3  0  1  0  1  4  5  2  3  3  1  20  2.0  4  0  1  0  3  2  4  5  2  4  1  22  2.2  5  1  0  2  0  3'  2  3  2  3  3  19  1.9  4 0.80 0.37  4 0.80 0.48  14 2.80 0.37  17 3.40 0.60  15 3.00 0.54  14 2.80 0.37  14 2.80 0.37  11 2.20 0.58  Total Mean S.E.  1 0.50 0.20  Kinoprene  8 1.60 0 . 60  application  to  f o u r•-day- •old p r e c o c e n e! t r e a t e d  102 " 20.40 0.67  10.2 2.04 0.07  insects  DAYS # INSECT  1  2  3  4  5  6  7  8  9  10  1  0  0  1  2  4  2  2  3  1  1  16  1.6  2  0  2  0  3  2  2  2  3  2  2  18  1.8  3  0  0  2  0  3  1  3  1  2  4  16  1.6  4  0  0  2  1  1  2  3  2  2  1  14  1.4  5  0  0  0  3  0  3  1  2  2  1  12  1.2  Total Mean S.E.  0 0 0  2 0.40 0.40  5 1.00 0.51  9 1.80 0.58  10 2.00 0.70  10 2.00 0.31  76 15.20 1.01  7.6 1.52 0.10  11 2.20 0 . 37  11 2.20 0 . 37  9 1.80 0.20  9 1.80 0.58  Total  Mean  106 Appendix 6 Taxonomic a u t h o r i t i e s  f o r s p e c i e s quoted i n t h e t h e s i s ;  COMMON NAME  L A T I N NAME  Green peach aphid Cabbage a p h i d Cowpea a p h i d Chrysanthemum a p h i d Potato aphid Vetch aphid Pea a p h i d B l a c k bean a p h i d Cockroach M i l k weed b u g Migratory locust Desert l o c u s t  Myzus p e r s i c a e Brevicoryne brassicae Aphis c r a c c i v o r a Macrosiphoniella sanborni Macrosiphum euphorbiae Megoura v i c i a e A c y r t h o s i p h o n pisum Aphis fabae P e r i p l a n e t a americana Oncopeltus fasciatus Locusta migratoria Schistoeerca gregaria  AUTHORITY (Sulz.) (L.) (Koch.) (Gill.) (Thos.) (Buckt.) (Harris) (Scop.) (L.) (Dall. ) (L.) (Forsk.)  

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