STOCK-SCION RELATIONSHIPS IN TOMATOES By ATMA SINGH A THESIS Submitted, to the State C o l l e g e in p a r t i a l School of G r a d u a t e S t u d i e s of Agriculture of M i c h i g a n and A p p l i e d S c i e n c e f u l f i l l m e n t o f the r e q u i r e m e n t s for the d e g r e e of D O C T O R OP P H I L O S O P H Y Department of H o r t i c u l t u r e 1949 ACKNOWLEDGMENTS The w r i t e r w i s h e s Dr. E. H. Lucas, State Col le ge , to e xp re ss h i s p r o f e s s or for his deep g r a t it ud e to of H o r t i c u l t u r e , i n s p i r i n g g u id an ce, he l p a nd e n c o u r a g e m e n t t h r o u g h o u t the Michigan unfailing course of these i nv es ti ga ti o ns . He also w i s h e s of the gu i d a n c e to e x t e n d his a p p r e c i a t i o n to m e m b e r s committee me n t of H o r t i c u l t u r e The v a l u a b l e and staff m e m b e r s for a d v i c e and suggestions coop er ati on . and h e l p g i v e n by Pr of ess or H. M. B r o w n , D e p a r t m e n t of F a r m Cr o p s , Colleg e, in the p r e p a r a t i o n g r a t e f u l l y a c k n o w l e d ge d . of the D e p a r t ­ M i c h i g a n State of the m a n u s c r i p t , are also CONTENTS Page INTRODUCTION 1 R E V I E W OF L I T E R A T U R E 3 GENERAL METHODS a ND M A T E R I A L S 11 Se l e c t i o n of P la nt M a t e r i a l 11 pro ce du re s 11 of G r a f t i n g D e t e r m i n a t i o n of To ta l So lu bl e and A s c o r b i c A c i d Solids , Acidity 13 EXPERIMENTATION 15 Stock-Scion Relationship With Reference Interaction Claimed by Lysenko Experiment to the (1947) 15 15 procedure 15 Results 19 and D i s c u s s i o n Experiment (1947-48) 26 Procedure 26 Results 30 and Discussion S t o c k - S c i o n R e l a t i o n s h i p W i t h R e f e r e n c e to the Syn th es is of A s c o r b i c A c i d and its R e l a t i o n to Total S o l u b l e S o l i d s a n d A c i d i t y 31 Procedure 32 Results 35 and D i s c u s s i o n S t o c k - S c i o n R e l a t i o n s h i p in C o n n e c t i o n W i t h Studi es o n r e c i p r o c a l G r a f t s of N o r m a l Plants and Plants M o d i f i e d T h r o u g h the A c t i o n o f 2 ,4 - d i c h l o r o p h e n o x y a c e t i c a c i d 44 Procedure 44' Results 45 and Discussion GENERa L DISCUSSION 50 SUMMARY AN D CONCLUSIONS ' 60 BIBLIOGRAPHY 63 INTRODUCTION The p r a c t i c e of g r a f t i n g is e m p l o y e d in g r e a t m e a s u r e be ca us e of the indivi dua l b e l i e f that st oc k a n d scion r e t a i n their ch ar ac te r i s t i c s . However, ences o f s to ck and trees a nd c o n s i d e r a b l e p r a c t i c a l us e has b e e n ma d e results blems s ci on are we l l the r e c i p r o c a l and e x p e r i e n c e s regarding gained e s t a b l i s h e d in m a n y the m e c h a n i s m of this the w o r k e r s qu an t i t a t i v e each p a r t ones consider these a n d s u g ge s t that is i n f l u e n c e d is th ro ug h of qu al iti es r e t a i n i n g its o w n i d e n t i t y years qualitative f r o m time reciprocal al ka loi ds acr os s herbaceous plants, but still scion i n t e r a c t i o n in fr ui t cised i n d i r e c t l y The r e su lt s influences changed nutrition. ef fe ct s It Ho we ver , is there each during recent the t r a n s l o c a t i o n of have been d e m o n s t r a t e d in the o b s e r v e d p h e n o m e n o n of trees as is m o s t l y e x p l a i n e d as stockexer­ t h r o u g h g r o w t h react io ns . concerning natur e of the scion, 1920), are not ge ne ra l to time. the only w a y in w h i c h such as the g ra ft un ion s, Pro­ at­ b e t w e e n s ci on and stock, intact. inf lu en ce s, of the interaction have ge n e r a l l y a s s u m e d that apart fro m n u t r i t i o n a l is no i n t e r c h a n g e fruit through experimentation. tr ac te d the a t t e n t i o n of h o r t i c u l t u r i s t s Most of influ­ the e f f e c t of the r e p o r t e d by D a n i e l (1897, s t o c k o n the 1904, 1911, in a c c o r d w i t h those n o t e d as h a v i n g m e t ac ce p t a n c e . H e has d e s c r i b e d m a n y cases the in which, -2- as a resu lt of gr a f t i n g , ch ara cte rs a s ho ot was p r o d u c e d c o m b i n i n g of s c i o n a n d sto ck a n d he v ar ia ti on s m a y be inherited. A considers s im ila r that such co n c e pt of "v ege tat ive h y b r i d i z a t i o n " has been held by certain scien­ tists in R u s s i a w h e r e a s t ro ng c o n t r o v e r s y has on since 19 o 0 b e t w e e n the " M e n d e l i a n " gen et ic is ts . The "non-Mendelian" b e e n h e a d e d by L y s e n k o . for Ly se nk o' s n e w on effects of g r a f ti ng . Hudson the data o f g r a f t h y b r i d i z a t i o n d i f f i c u l t y of i n t e r p r e t a t i o n . a r e p e t i t i o n of strictly scion is not controversial physiology selected. co nd iti ons . for genetic c o m p e l l i n g but It was w i t h experimental is f o r m e d by and R i c h e n s after carefully They emphasize basis evidence (194b) in every detail, has in analyzing m e n t i o n the the u r g e n t n e e d some of L y s e n k o 1s e x p e r i m e n t s u n d e r controlled that "the e v i d e n c e school of g e n e t i c s theories their r e v i e w of the n e w the ories, for an d " n o n - M e n d e l i a n " The p r i n c i p a l g e n e ti c al been going the points that this In summarizing the y interaction between state stock and sug ges ti ve ." a i m of a n a l y z i n g some o f th es e h i g h l y in the f i e l d of g e n e t i c s and p l a n t p r o b l e m of s t o c k - s c i o n i n t e r a c t i o n was R E V I E W OP L I T E R A T U R E Prior to 1946, papers h a v e se ve r a l h i s t o r i c a l a p p e a r e d In v a r i o u s c o n t r o v e r s y in the U.S .S .R . pi na ss e 1941, H a l d a n e prehensive (Sax 1944, 1940, M a t h e r available l a t i o n of L y s e n k o ' s b o o k (1946) j o ur na ls on the gen et ic s Pinc us 1942). 1940, The i n f o r m a t i o n in E n g l i s h o n L y s e n k o ' s though t was m a d e w h i c h was and. t h e o r e t i c a l first p u b l i s h e d provide in the S o vi et additional union" Es- f ir st coinscho ol of in 1946 b y D o b z h a n s k y 's t r a n s ­ "Heredity a n d its V a r i a b i l i t y " , in 194b. H u d s o n and R i ch en s i n f o r m a t i o n on "The New G e n e t i c s and d i s c u s s the results o b t a i n e d by Russian scientists. The m o s t by Lys enk o, a r o u n d the i m p o r t a n t a nd r a d i c a l l y as r e v e a l e d by D o b z h a n s k y ' s id e a of d i t y of o r g a n i s m s vi ron men t. body He inheritance considers to r e q u i r e definite according accepted, conditions to M e n d e l i a n heredity heredity regards " h e r e d i t y as definitely t r a n s la ti on , controlled by the the e n ­ for its to va ri o u s life, its d e v e ­ co nd it io ns ". How­ t h e o r y w h i c h h as b e e n g e n e r a l l y characters This v i e w of of an o r g a n i s m are the res ul t of an i n t e r a c t i o n b e t w e e n its H e r e d i t a r y ge n o t y p e the e n v i r o n m e n t Here­ the p r o p e r t y of a l i v i n g is c o n t r o l l e d b y the genes. that cen te r of a c q u i r e d ch ar act ers . a c c o r d i n g to h i m is lo pm en t a n d to r e a c t ever, d i f f e r e n t v ie ws h e l d in w h i c h it d e v e lo ps . Altering and the env iron- -4- ment m a y change not in herited; these this characters is what of a c q u i r e d c h a r ac te rs. is but genetics (1947) A permanent hereditary considers g r e w o u t of h i s pr ob a b l y his success environmental classical work genotype was could be not jecti on of o r t h o d o x g e n e t i c s of the Da vi e s and its followers evidence w h i c h they Fyfe progress gests (1947) whi le r e p l a c e m e n t by d i s c o v e r e d new of re­ the Dolgusin experimental been adequately nea rly a h a l f - c e n t u r y (1946), that In any of s c i e n t i f i c sug­ ex­ reviewing Lysenko's "Lysenko may case his re ce n t of the tested. a n d its v a r i a b i l i t y , states facts. theories judgment.on Lysenko's and G o l d s c h m i d t btern Prezent, a n d d i s a g r e e i n g w i t h his co nc e p t i o n of h e r e d i t y mai n points, thatk'tne m e n t i o n i n g L y s e n k o ’s .i g n o r a n c e they h a v e (194b) in submit" . the w i t h h o l d i n g Stern ch ang es is qu it e u n j u s t i f i e d b y the in ge n e t i c s pe riments u n t i l and environment states form o f L a m a r c k i s m a d v o c a t e d b y L y se nk o, and their the d i s t i n c ­ inheritable p r o d u c e d by m o d i f i c a t i o n s which the o r g a n i s m d e v e l o p e d . that t h r o u g h an between phenotype a n d that in on v e r n a l i z a t i o n , l e d h i m to co nc lu de co rrect, interest the p h e n o t y p e tion d r a w n by the M e n d e l i a n g r o u p change the g e n ot yp e. that L y s e n k o ' s in a l t e r i n g change Is called-non-inheritance can be a c c o m p l i s h e d o n l y by a l t e r i n g Davies such a cha ng e criticize the or m a y n ot ha v e attempt to i g n o r e achievements seems to -5 have r e s u l t e d in r e p l a c i n g r a t i o n a l and va gu e by h u n c h e s generalities". Darlington (1947), Soviet Sci en ce ", quotes dity f r o m D o b z h a n s k y * s of ar gu m e n t idea as in si g h ts in his "A R e v o l u t i o n L y s e n k o ’s views He the line i n t e r p r e t s L y s e n k o ’s m a i n " H e r e d i t y is d e ve lo pm en t. can change d e v e l o p m e nt . in wi t h r e g a r d to h e r e ­ t r a n s l a t i o n a n d c ri ti ci ze s f o l l o w e d by him. follows: articl e Therefore the The e n v i r o n m e n t e n vi r o n m e n t can change heredity". Ashby (1946) lo wed b y Ly s e n k o finds that the experimental and o t h e r w o rk er s that the plant m a t e r i a l of h i s is h e t e r o z y g o u s , emp lo ye d are i n s u f f i c i e n t and the t e c hn iq ue school is poor, the p h y s i c a l statistical f ol ­ controls an al ys is of data is lac kin g. In v i e w of the d i s a g r e e m e n t s and c o nc lu si on s views of the R u s s i a n s in ge n e t i c s tion of at least in the r e p o r t e d observations a n d the it s e e m e d d e s i r a b l e the m o s t generally accepted to at t e m pt s t r i k i n g of the R u s s i a n ex pe ri me nt s. m e n t i o n e d ear lier, L y s e n k o h as p l a c e d special gra fti ng exp eri me nt s, of w h i c h he c i t e d se ver al as support of his fore, to revie w theories. the earlier w o r k in this the r e p e t i ­ It w o u l d not be "graft-hybrid" co nn e c t i o n . emp ha si s re sul ts out of place, c o n c e p t and some of on in th er e­ the Weiss various states (1950) pr o b l e m s and Jones (1954) have r e v i e w e d the r e l a t i n g to g r a f t h y b r i d i z a t i o n . that ''the nature of 'graft hybrids' has cei ve d so d i f f e r e n t l y b y d i f f e r e n t wo rk e r s, with su c h a l a c k know of pr eci sio n, ex act ly w h a t that it Jones been con­ and o f t e n is i m p o ss ib le to is i m p l i e d i n any g i v e n case w i t h o u t fu rt he r d e f i n it io n. " The m o s t of a tr u e graft h y b r i d is, clearly that expressed version it re su lt s f r o m the fusion of s o m a t i c n u c l e i of s ci on and stock in a m a n n e r a n a l o g o u s to tha.t in w h i c h rise to fusion b e t w e e n ga m e t e s a seed hybrid.. of a g r a f t hy br i d , and s t o c k w i t h o u t rather p e c u l i a r Jones hyb rids, ferent that th ere has is no Daniel s ci on is states j u s t i f i c a t i o n f o r re g a r d i n g as by nuclear that cells. "the o c c u r r e n c e is sped are dif­ the r e s u l t of f u s i o n of eith er C o m m e n t i n g on the h y p o t h e ­ of nuclear fusion dissimilar vegetative of gr aft h y b r i d is n ot sach f u s i o n sug­ fundamentally strains the g e n e t i c a l l y to the p r o d u c t i o n of D a ni el who that all or v e g e t a t i v e possible; q u a li ti es o f co ns ide rs sex-cells between conception of n u c l e a r fusion, and c h i ma er as of tv/o plant Jones the s t at ed the views the u n i o n sis, it co m b i n e s the o c c u r r e n c e graft h y b r i d s types. some wo rkers' give and h a r d to exp lai n. (1954) ges te d tha t However, or se x- ce ll s cells leading theoretically certainly exceptional im­ and s ti ll -7- awaits dem on st ra t io n' 1. Weiss (19d0) is also i n c l i n e d to the o p i n i o n that h e r e d i t a r y interaction between and sc ion of g r a f ts h a s not yet been demonstrated. Hu d s o n and Richens Ly se nko 's (1946), graft hybridization theory of v e g e t a t i v e wh il e tracing theory, rapprochement the s t o c k the b a s i s mention Michurin*s and Timirjazev's D a r ­ w i n i a n n o t i o n of g r a f t h y b r i d i g a t i o n . According urin, the g e n e t i c a l n a t u r e s elements gr aft s h o u l d a p p r o a c h ea ch oth er . act io n of g r a f t i n g changes. His is sufficient tant b a s i s the recent of Lysenko's ting of tom at o varieties to h a v e experiments genetical been supported by w h i c h form t h e o r ie s, habits gr owt h. car pe ls , These characters may and v e r y y o u n g p l a n t s be as p ap er w h i c h s u p p o r t s L y s e n k o ' s Jastr eb (1941). and Ric he ns . grafting the leaf scions. views variety Albino (stock). as a nd have shown transmitted from stock is Especially interesting-are va r iety, M e x i c a n 3 55 impor­ shape, experiments It has b e e n t h o r o u g h l y tom at o the involved graf­ scion u n d e r p r e s c r i b e d c o n d i t i o n s u s i n g m a t u r e stocks Ric he ns . d i f f e r i n g in s u c h c h a r a c t e r s n u m b e r of that all th ese of the that the m e r e c i t e d b y H u d s o n and fruit size or color, of v e g e t a t i v e assumed to M i c h ­ to b r i n g a b o u t h e r e d i t a r y assumption appears a n u m b e r of R u s s i a n pa pe rs M o s t of o f the two He of The m o s t for important that o f A v a k i a n and r e v i e w e d by H u d s o n the r e s u l t s (scion)onto It has plants to the of red b e e n c l a i m e d that the w h i t e s c i o n sh oo t bore frui ts of v a r i o u s ranging f r o m r e d to pale y e l l o w . and Fo g e n e r a t i o n s colors Further records s h o w e d a g r a d a t i o n of colors. of F^ The (Zj reciprocal effect of scion o n stock was also s ho wn b y A v a k i a n and J a s t r e b b y u s i n g A l b i n o v a r i e t y as and M e x i c a n 3 5 5 characteris­ tics of as scion. the s c i o n we re Results of several In this i m p a r t e d to the other Richens. .cereals as d i s c u s s e d above, In a d d i t i o n , like w h e a t, ch ara cte rs of b o t h Outside support to some s t o c k and Daniel's principally on fruit trees but his some o f scion, 1904, findings Wettstein and Pirschle def e c t i v e petunia to n o r m a l plants that some apple varieties, are is f o u n d in As m e n t i o n e d 1980) ex pe r i m e n t s , spp., d i d show induced by mere grafting c on fi rm ed . (1958) b y g r a ft in g. b el ow given. ev i d e n c e 1911, found could transmit leaf ties p r o d u c e d f r u i t s the m o r p h o l o g i c a l concept. c an be have not b e e n d.S .S. R. on g r a f t i n g of or Helianthus that h e r i t a b l e m o d i f i c a t i o n s the are p r e s e n t e d by H u d s o n a n d v e r y l i tt le (1897, in plants- and d e p i c t i n g s i m i ­ o f the g r a f t h y b r i d i z a t i o n pre vi ou sl y , stock. experiments combine the U . S . S . R . the investigations involving grafting of herbaceous lar ch ang es case stock that a c h l o r o p h y l l a n d shoot Heinicke (1927, ch ar ac te rs 1936) noticed- w h e n top w o r k e d w i t h o t h e r v a r i e ­ the graft u n i o n s displaying -9- ch ara ct er s o f the scions. r in g i n g the b r a n c h e s the scion effect . (1955) ap ple ha ve that the s c i o n had, problems stock Wa rn e po s s i b l e did not c h e m i ca l sci on s The co nc l u d e c h a ra c te rs that many o f examined. gra ft s f o u n d no fcnd K e r k i s chemi­ the o ut ­ on the Hofmann (1927) through b e a n and w h i t e Wavy d i d n o t obser ve the m o v e m e n t of r e d beet a n d sugar t r a n s f e r of inu li n f r o m ar t i ­ stocks. c e r t a i n a l k a l o i d s acr os s b e e n d e m o n s t r a t e d from time sci on s explained of a n t h o c y a n i n pigment (1902) sunflower of between an i n v e s t i g a t i o n of the could n o t be s h o w e d that a t r o p i n e Belladonna (1941) (1926) of the rootstock. b e t w e e n r o o t s t o c k effect a n d transfer transfer u n i o n has (1894) ha v e m a d e in r e c i p r o c a l to an a p pr e c i a b l e relationship of r e d d i s h pu rp l e R e f u g e e of red p i g m e n t choke instances, T h e se w o r k e r s f i n d the Daniel T h e y ha v e p o i n t e d out be e n i n v e s t i g a t e d by m a n y workers. (1955) Similarly Biffen beet. a n d Tukey a n d Erase g r o w t h and c h ar ac ter r o o t s t o c k ef fec ts graft u n i o n s bean. in some and s ci on ha v e of the (1950), rootsto ck s. relationships sta n d i n g the fruits h e l p e d i n t e ns if y r e g a r d i n g the che mi ca l cal fac tor s. basis the and W a l l a c e and at l e n g t h w i t h the effects of various on their influence upon the below Swarbrick de al t scions R e m o v i n g st oc k leaves pa s s e s into pot at o graft Daniel r e a d i l y from A t ropa stocks. and P i g u l e v s k a y a to time. the Kraevoi (1941) a n d .Nechaev found th a t atro- -lO- pine was translocated from Datura stramonium rootstocks L v cop er si c um e s c u l e n t u m scions. that n i c o t i n e p r o d u c e d across nicotin e into while s pe ci es (which la ck s are of ta bacum, it is u s e d as the L y s e n k o has roots tomato the and a p r e c i p i t a b l e b a s e d his the o ri es the since accessible, to r e p e a t scion in a graft w i t h amount some ve ry firmly o n results His claims t he to be meager. of his solids, we r e det er mi ne d. grafts of n o r m a l as and h is total some In addition, plants deemed f o l l o w e r s ’ tomato o u t l i n e d in physiological an d u n g r a f t e d plants acidity and as co rb ic I n ord er characters -- t o t a l acid c o n t e n t a s t u d y was m a d e on a n d pl an t s xaodified t h r o u g h of 2 , 4 - d i c b l o r o p h e n o x y a c e t i c acid. some­ as far It wa s c l os el y as possible. stu di es , of g r a f t e d are e x p e r i m e n t a l evidence, appears the H u s s i a n p u b l i c a t i o n s soluble in the in t a b a c u m w h e n gra ft in g e x p e r i m e n t s , f o l l o w i n g the tec hn iq ue s of the fru it s and s c i o n stock. what surprising, to sub st an ti at e of tabacum x of anab as in e a p p e a r s experiments. de si ra bl e the i n h e r i t a n c e an d o b s e r v e d that w h e n t a b a c u m is u s e d as amount shown Kuzmenko and i n t e r a c t i o n of s to ck rep or te d f r o m g r a f t i n g as it is has is t r a n s p o r t e d scions. w o r k i n g on gra fted, a nab as ine ) a considerable (1942) co n t e n t of H i c o t i a n a g l au c a h y b r i d s w h e n these glauca, (1940), and a n a b a s i n e H i c ot ia na leaves in tobacco the gr af t u n i o n s T ik hv in sk ay a Dawson to -- reciprocal the action -11- GENERAL METHODS Selection of Plant fering in color, red), Stokesdale I ta li an R e d Pear (small, red, The seeds size a n d n u m b e r of l o c u l e s 1946. T h e s e w er e; (small, pear-shaped), red, (large, were Incidentally, the which were rec o r d e d v a r i e t i e s , (1946) one for high ascorbic l e c t i o n in 1947 Procedures for in t o m a t o e s the a b ov e line d e v e l o p e d b y Y e a g e r a n d P u r i n t o n aci d co ntent, of Grafting i l l us tr a t e d in f i g u r e I w a s terminal seed com­ s e l e c t e d b y Yiiilson a n d W i t h n e r 1 9 4 o . ’ In a d d i t i o n to studies of the i n v e s t i g a t i o n s white)„ s e l e c t e d wer e (1946) w h o s e w o r k on s t o c k - s c i o n r e l a t i o n s h i p s was r e p o r t e d in D e c e m b e r , yellow), g u a r a n t e e d for some of the v a r i e t i e s those (large, (large, obtained from commercial same as of f ru it Italian Red Plum p l u m - s h a p e d ) , and W h i t e B e a u t y varieties, dif­ John Baer Golden Queen of th ese found to be - Tomato varieties (l arg e,redj, purity of st rains, panies. ND M a T E R I a LS Material shape, we r e s e l e c t e d in fall, a on a s c o r b i c wa s a d d e d to acid and mostly p o r t i o n of the p l a n t in the in the grafting f i rs t s e c o n d year. (to be u s e d col­ s y n t h e s i s .# - The m e t h o d o f spli ce used entirely the as stock), year The in- *The seed of t o m a t o w i t h h i g h a s c o r b i c a c i d c o n t e n t was o b ­ tai ne d by Dr. S. H. W i t t w er , M i c h i g a n S t a t e C o l l e g e , h a s t Lansing, M i c h i g a n , t h r o u g h the c o u r t e s y of Dr. A. P. Ye ag er , U n i v e r s i t y of New H a m p s h i r e , D u rh a m , N e w H a m p s h i r e . Fig. I Technique of G r a f t i n g C o n s e c u t i v e steps in the p r o c e d u r e whe n s c io ns from t h r e e - w e e k o ld p l a n t s w e r e g r a f t e d onto s i x - w e e k old s t o c k s „ Figure I Pig. II Technique of G r a f t i n g Co ns e c u t i v e s t e p s in the p r o c e d u r e w h e n scions with two c o t y l e d o n s an d two small le a v e s were g r a f t e d onto s i x - w e e k old s t o c k s „ Figure II -12- cludi ng the apex and y o u n g leaves, obliq ue cut. was a s i m i l a r cut was m a d e r e m o v e d v.-ith an on the p o r t i o n of the c o m p a r a t i v e l y y o u n g e r p l a n t scion) . The cut s u r f a c e s tog e t h e r a n d then h e l d narrow strip of wet at mo s p h e r e in place b y raffia until The w r a p p i n g them w i t h a fiber as sho wn in fig ur e I (1-5). the p l a n t s we r e p l a c e d in the of c h a m b e r s , six 4 - i n c h pobs. each of which h e l d t h ir ty - pl an t s were k e p t in the the grafts h ad b e g u n to u n i t e seven to ten days. hi gh b y In oc ca si on al ly the me a n t i m e graft u n i o n s . non-sjjecific Pr op er ta ken had t a k e n p lac e and w e r e of the gre en hou se . of p r o p e r uni on, for a n o t h e r few days. fia f i b e r was out If the re m o v e d H o w e v e r, fu ngi w h i c h d e s t r o y e d the of the c h a m b e r s t r a n s f e r r e d to scions the damage. w h e n some h e a l i n g the s h a d e d s e c t i o n showed dropping owing to l a c k p l a n t s were a g a i n p l a c e d in the W h e n the u n i o n was ca r e fu ll y b y too p r o v e d c o nd uc ive a e r a t i o n h e l p e d to redu ce The p l a n t s we re took the h u m i d i t y was ke p t s pr in k li ng w i t h water. d e v el op me nt of chambers wh ic h u s u a l l y high a h u m i d i t y and t e m p e r a t u r e a l w a y s to the (to be u s e d as of stocic and s c i o n w e r e b r o u g h t I m m e d i a t e l y after g r a f t i n g , humid t e r m in al cu tt i n g co mp l e t e d , it into chambers the raf ­ small strips. T h e me t h o d , l i m i t e d e x t e nt order to i l l u s t r a t e d in f i g u r e in the s e c o n d year o f II, was u s e d to a the i n v e s t i g a t i o n s , in study the s t o c k - s c i o n i n t e r a c t i o n wh e n v e r y y o u n g -15- se ed l in g s w e r e u s e d as plants be u s e d as to split o p e n b y cotyl of leaves, was sert ed i n t o of raff ia seedling, slit were It took a b o u t two w e e k s that t h e the plants wer e analyzed were weighed at one time. separately fruits. ti ma ti on o f soluble The sample dry the use meant was of in f i gu re II steps One small strip (2-5), taken, to se cu re d i s c u s s e d i n the f i r s t m e t h o d . to form sufficiently from the m o i s t golids, ripe fo u r to e i g h t fruits samples fro m eac h p l a n t of 1 0 0 g ra ms s a m p l e was solids and chamber. .acidity and the l o n g i t u d i n a l l y for to ta l each cut s e c t o r s u s e d fo r acidity Th e the and the es­ other aliquot was so l i d s filtered through a tested .acidity wa s of the and a cidity of distilled water and tnen filtrate a refractometer. t r a t i n g a 2 0 .ml. soluble c r u s h e d in IG u ml. of a W a r i n g B l e n d o r fil te r p a p e r . b y means two acid determination. determination, by and a Further Two fr o m The h y p o - stock. determinations, fresh and fully total shown be r e m o v e d these of' side o n l y a n d i n ­ the for the u n i o n of r e p r e s e n t a t i v e for a s c o r b i c as in of T o t a l S o l u b l e A s c o r b i c .noid - For representative same could Determination cut o n o n e in pla ce. a good u n i o n , grafting. cotyledons as portions d e c a p i t a t e d and later already made seedling terminal c l ef t w i t h two f i b e r was w r a p p e d , to ho l d t h e so if for g i v e n an o b l i q u e the The stoc ks w e r e a cut as the s ci o ns . filtrate for t ot al solids determined by against ti­ standard .-14sod iu m h y d r o x i d e u s i n g p h e n o l p h t h a l e i n as the i n d i c a t o r . The m e t h o d of c r u s h i n g tity of water, and t h e n t r a n s f e r r i n g to a v o l u m e t r i c adopted , fl as k because of u n k n o w n the fruit in a l i m i t e d q u a n ­ the crushed material and m a k i n g up to volume, was n o t d u r i n g the p r oc es s of bl e n d in g, and u n e q u a l cess of m a k i n g u p to quantities intr od uct io n of a i r h i n d e r e d in the p r o ­ the fi na l volum e* .ascorbic a c i d d e t e r m i n a t i o n s were made a f t e r cedure of Lucas (1944). was p l a c e d in 5 0 0 ml. a Waring Blendor fi l t e r e d of A f r e s h 1 0 0 - g r a m s a m p l e of f r u i t 2 per cent m e t a p h o s p h o r i c a n d cr ushed. The crushed material t h r o u g h d r y f i l t e r paper. was t i t r a t e d a g a i n s t indophenol An aliquot a c i d in was of f i l t r a t e s t a n d a r d so di u m 2 ,6 - d i c h l o r o b e n z e n o n e - s o l u t i o n an d the a m o u n t of a s c o r b i c p r e s s e d as m i l l i g r a m s the p r o ­ per 100 grams of ac i d wa s fre sh fruit. ex ­ EXPERIMENTATION Stock-Scion Relationship with Reference a c t i o n C l a i m e d by L y s e n k o As has b e e n stated i n the ticists hav e b e e n ra th e r f oll owe r's ideas. to an a c c e p t a n c e Many "Review of the Lysenko's to c o n t r i b u t e a cr it ic a l ting e x p e r i m e n t s (1941) by school. them as their re p o r t s p e r m i t t e d . it prov id es , of the The p r e s e n t e x a m i n a t i o n of with tomatoes repeating and h i s c o n c l u s i o n s d r a w n by Ly se n k o , the R u s s i a n gene­ although strictly opposed st re s s e d the n e e d for a r e p e t i t i o n of so m e d e s c r i b e d by the I n t e r - of L i t e r a t u r e " , cr i t i c a l about of there, to the r e s u l t s o f g r a f ­ and J a s t r e b th e d e t a i l s Their work was a c c o r d i n g to L y s e n k o , experiments study attempted claimed b y Avakian c l o s e l y as have g i v e n in s i n g l e d out b e c a u s e the m o s t n o t a b l e support for his h y p o t h e s e s . Experiment Procedure - Seeds of (1947) all the v a r i e t i e s tioned 'were sown in s e p a r a t e January, 1947. ten days old, Uniform were seedlings of Ap ri l , procal gr a f t s greenhouse during of a l l v a r i e t i e s , t r a n s p l a n t e d into 4 - i n c h n e w plants w e r e g r a f t e d d u r i n g two weeks fl at s in t h e previously m e n ­ when pots. the l a s t w e e k of M a r c h they were 50 to w e r e ma d e b e t w e e n J o h n B a e r S 5 cm. eight to Th e and f i r s t tall. Reci­ and W h i t e B e a u t y -16- an d b e t w e e n I t a l i a n P e a r a n d White Beauty. the rest of the v ar ie t i e s , an d I t a l i a n P l u m, as the were scion in St o k e s d a l e , they w e r e u s e d as each case. grown later viz., In the case of* stocks w i t h White B e a u t y The pl a nt s so that v e r y small G o l d e n Qu een to be u s e d as s c i o n s te rm i n a l parts w i t h o u t any f l o w e r p r i m o r d i a c o u l d be o b t a i n e d and g r a f t e d onto o l d plants. Usually t h r e e - w e e k old seedlings, were u s e d for o b t a i n i n g t e r m i n a l p or ti on s Th e n um be rs shown in table Table 1. of total a n d su cc es sf ul 8 to 10 cm. t a l l , to serve as s c i o n s . graf ts in 1947 a r e 1. N u m b e r s of p l a n t s of v a r i e t i e s d i f f e r i n g in f r u i t cha ra ct er s, w h i c h were g r a f t e d and w h i c h s u r v i v e d d u r i n g 1947. Graft combinations Stock Scion Fruit character differences Number Number C o l o r Shape Size N o . o f grafted survived lo o u l e s John Baer White Beauty 56 51 X - - “ Wh it e B e a u t y J o h n Baer 55 28 X - - - Stokesdale White Beauty 56 51 X - - - I t a l i a n Pea r White Beauty 56 55 X X X X White Beauty I t a l i a n pe a r 56 3U X X X X Italian- Pl u m Vi/hite B e a u t y 06 55 X X X X Golden W h i t e Beauty 56 54 X - — — 252 224 Queen Total x indicates a difference i n the c h a r a c t e r b e t w e e n two v a r i e t i e s . -17- Most one from of the graft ed p l a n t s the l im i t e d to stock and o n e one shoot lateral b r a n c h e s m o v a l of t he ro w s The 5 feet apart, 5 feet. staked the scion, scion. by Th e were the and the f o l l o w i n g some w e r e development were also of re­ removed' t r a n s p l a n t e d into distance Soon a f t e r but shoots, regular and frequent f l o w e r buds 6 , w h e n the p l a n t s also b e i n g were from suppressed a x i l l a r y buds. un ti l J u n e fi eld in was from w e r e t r a i n e d to two the between plants transplanting, experimental the p l a n t s tre at me nt s started; (a) O n l y one shoot of and bear frui t. piece of stem. were r e m o v e d . The ing the i n f l u e n c e (b) Two and the three s m a l l one f r o m The al lo w e d to leaves They fruit. The leaves on the tact but the flower The the i n f l u e n c e fruit. the s et an d a s m a l l the g r a f t u n i o n s scion fruits stock the of s to ck l e a v e s , apex, two were removed this but as treatment in the scion. from or left had f r o m the s t o c k w e r e were study­ defoliated On ly a f t e r two c l u s t e r s shoot in a n d the o t h e r sh oo t w a s ke pt shoot for f r u i t of the f r u i t on it. also r e m o v e d clusters on the scion protecting scion. appeared. were only roots d e v e l o p i n g b el ow of the r o o t s t o c k trained. flowers allovted to dev elo p, treatment provided conditions shoots, the s c i o n w e r e s c i o n was s toc k p r o v i d e d All buds This t he on the set left in­ soon as t h e y developed under absence of s t o c k -18- (c) This and p r o v i d e d t r e a t m e n t was conditions fruit u n d e r the the o p p o s i t e of for s t u d y i n g the influence treatment development of sc ion l e av e s b u t (b) of s t o c k in the a b s e n c e of s c i o n fruit. (d) fruit. B o t h s t o c k and s c i o n shoots ho le av e s were re mov ed . were This allowed treatment to bear s e r v e d as a g r a f t e d con tro l. T he graft number combinations plants of pu rp os e is s h o w n in table ea ch v a r i e t y we r e ste m a n d Out of a l l o w e d to six p l a n t s 2. were these, si x pla nt s set f r u i t t r a i n e d to shoo t, m i t t e d to set leaves were (t re a t m e n t fr om d i f f e r e n t I n ad di ti on , gr ow n on th eir o w n roots shoots. allowed left (f))c intact twe lv e for the and to serve as ’were t r a i n e d to a sin gl e (treatment two d e f o l i a t e d a n d the fr ui t ot her treatment of c o m p a r i s o n of f r u i t c h ar ac te rs , co nt ro ls . was of plants u n d e r e a c h (e)>. One of The these to set o n it; but no remaining shoots on the fr uit was per­ -19- Table 2. N u m b e r of plants Gr aft combination Sto ck of di f f e re nt gr^ ft Number Scion co mb ina tio ns of graft sy mbionts u n d e r various tr eat men ts (a) (b ) (c) (d ) To ta l J o h n Bae r White B ea u t y 5 8 6 4 25* W hi te Jo h n Baer 4 10 8 .6 28 Stokesdale White Beau ty 5 10 10 6 31 I t a l i a n Pear W hit e B e au ty 6 12 11 6 35 'White B e a u t y I t a l i a n Pear 5 12 8 5 oO Italian Plum 'White B e a u t y 6 12 n 6 35 Golden Queen W h i t e Bea ut y 6 12 10 6 34 37 76 64 89 216 Beauty To tal f u l l y g r a f t e d pl an t s Results fru its normal. d i e d s o o n after t r a n s p l a n t i n g in the field. and D i s c u s s i o n - T he on d e f o l i a t e d shoots Obviously, the No plants , ployed. d i d not ch ang es we r e of treatm ent s fruit elaborated food materials in vi sib le de ve l o p m e n t and m a t u r i t y of (b) and fro m the other ch ar ac t e r s of fru it s e x p e r im en ta l of lo cu le s of fruit, in color, dr e w symbiont* shape, from g r a f t e d tr ea tme nt D e f o l i a t i o n and d e f l o r a t i o n of e i t h e r in change s were o n the d e f o l i a t e d shoots p r o d u c e d u n d e r an y re su l t (c) em­ stock or scion size or n u m b e r w h i c h c o u l d be a t t r i b u t e d to the treatments. The plants u n d e r stock r o o t s mal f r u i t s grafted t r e a t me nt alone o n of the scion f r u i t was s c i o n var ie ty . control p l a n t s results are by W i l s o n a n d Avakian and Jastreb si mi l a r varie tie s to tne n u m b e r o f plants in e a c h treatment sions. In the p r e s e n t plants under significance Of the was to the fruit but fail re­ to c o n f i r m the symbi ont s r e p o r t e d by and Wi t h n e r investigations (1946) used them any s ol id conclu­ the num be r o f gr af t e d sufficient to give a high r es ul ts . T h e v a r i e t y W h i t e B e a u t y was n o t of fruit to the ripening, of f r u i t trees, v a r i e t y Whi te in gra ft s Ho we ve r, effects b e t w e e n e a r l y and basis. It of h a v e b e e n r e p o r t e d q u i t e o ft en Beauty was in fruit b e a r i n g . ap­ time of m a t u r i t y the o t h e r va ri e t i e s . and e x p l a i n e d on n u t r i t i o n a l p r ol if ic s ym ­ six v a r i e t i e s u s e d W h i t e B e a u t y and G o l d e n ^ u e e n late v a r i e t i e s The Each g r a f t g r a f t e d and em pl o y e d b y influenced with regard ti m e of developed normally too l i m i t e d to draw w h e n grafted o n on the the f r u i t s in the p r e s e n t i n v e s t i g a t i o n . each t r e a t m e n t was were l a t e - r i p e n i n g . preciably Wilson those u s e d produced nor­ a g r e e m e n t with t h o s e of gr af t (1941). of its o w n variety. Vi/ithner (1946) in fruit c h a r a c t e r s H ow e v e r , (d)) of in p e r f e c t changes studied, characte rs. fruit c h a r a c t e r i s t i c These ported in the i n f l u e n c e Similarly, (treatment and s h o w e d no m o d i f i c a t i o n s biont b o r e (a), w h e r e f o u n d vigorous and v e r y was h i g h l y c om pa ti bl e wi th -21- the o t h e r va r i e t i e s empl oye d, and no d i f f i c u l t y was p e r i e n c e d 'while g r a f t i n g it wi t h other* va rieties, e v i d e n c e d by the n u m b e r of s u c c e s s f u l In con tr as t, W i l s o n a nd W i t h n e r a p p r e ci ab le in dicates analys is varieties under of fruit study, solub le Hence a study o f the fruit solids, ti tratable was u n d e r t a k e n in the ficant of g r a f t o, 4 a n d s y mb io nt s, The did. n o t an­ to ta l of fruits s o lu bl e from show the cont ro ls . 1947. each r e s u l t s of 5. acid c o n t e n t when compared with treat­ of S e p t e m b e r , The acid. reciprocal of f r u i t from f o u r pl a n t s of and a s c o r b i c differences, of the under different a n a l y z e d s e p a ra te ly . treat me nt s the B e a u t y w i t h r e g a r d to first w e e k are p r e s e n t e d i n ta bl e s acidi ty of all a c i d i t y and a s c o r b i c two v a r i e t i e s , samples plants composition, ments, were gr a f t s success. a n d Wh it e of on l y these Representative However, on h e t e r o p l a s t i c f r o m c o nt ro l grafts different fi n d s h o w e d that m a x i m u m d i f f e r e n c e s we r e total solids, 1. that even grafts b e t w e e n u n r e l a t e d plan ts e x h i b i t e d b e t w e e n J o h n Ba e r alyses (194b) , who d i d not (1948) can be m a d e w i t h c o n s i d e r a b l e treatment, is in t a b l e Wh it e a p o o r g r a f t i n g variety. the r e c e n t ‘work of N i c k e l l The as d i f f e r e n c e s b e t w e e n the v i g o r of v a r i e t i e s used, c o n s i d e r e d Cryst al clear ly grafts ex­ signi­ -22- Table 3. Effect solids v O6 of d i f f e r e n t t re at me nt s on the c o n te nt of fruits, V J- J. a.wu-u p 1947. T r e a t ­ Fr uit Graft combination ments* analy­ zed total F e r cent total solub le fruits of 4 i nd iv id ua l soluble solids of pla nt s + Stock— Scion 1 2 5 4 M e a n S .E . JB WB (a) WB 4.4 5,6 4.0 5.8 5.9 .17 JB WB (b) WB 3 c8 4.0 4.o 4.0 4.1 .20 JB WB (c) JB 5.2 6.0 5.4 5.6 5.5 .17 — JB 6 .2 5.6 6.0 5.8 5.9 .15 JB WB ""-WB 4.6 4.0 4.0 0.8 (d)C. 4.1 .17 JB (e) JB 5.4 5.8 6.2 5.6 5.7 .17 JB (f) JB 5.2 5.0 6 „0 5.4 5.4 .22 WB JB (a) JB 5.8 5.0 6 .2 5.6 5.6 .2 5 IB JB (b) JB 5.2 4.8 6 .0 5.4 5.5 .25 ‘ WB JB (c) WB 4 .4 4.0 5.4 0.8 5.9 .21 - WE 4.6 4.4 4 .0 3.8 4. 2 .18 WB JB '•‘‘JB 5. 4 5.6 6.0 5.0 (d)C 5.5 .21 WB - (e) 7© 4.-4 5.6 4.2 0.8 4 .0 .18 WB (f) WB 5.6 4.0 4. 0 o.4 5.7 .15 A' . VJ + ■G >\ 1 A n 3, X -K-For tables JB and WB res pec tiv ely . c; A o > 4 and. 5. i nd ic at e J o h n Ba e r end W h i t e B e a u t y v a r i e t i e s Treatments: (a) (b) One sho ot fr o m scion, w i t h fr u it s and leaves . Two shoots; one f r o m scion, de fo li ate d, w i t h fruits; one from s t o c k d e f l o r a t e d , with leaves, (c) Two shoots; one fr o m scion, de fl ora ted , w i t h leaves; o n e fro m stock; d e f o l i a t e d , wi t h fruits, (d) Two shoots; one fr o m s c i o n and one from stock; b o t h w i t h leaves a nd fruits. (e) One shoot wi th fruits a n d leav es (un gr af te d control). (f) Two shoots; one d e f o l i a t e d ;,ith fruits, the o t h e r d e f l o r ­ ate! w i t h leave s (un gr af te d, control). -26- Table 4. E f f e c t of d i f f e r e n t t r e a t m e n t s on the ti tr at ab le a c i d c o n t e n t of f r ui ts , 1947»_____ . _________________ _ Graft c o m bi na ti on Treat­ ments* Fruit A c i d i t y in terms of ml. of 0.1 N NaOH a n a l y ­ u s e d to t i tr a t e a 20 ml . a l i q u o t zed Stock - Sc ion 1 2 6 4 M e a n + S.E. JB WB (a) WB b .4 7.0 6.8 5.7 6.5 .29 JB WB (b) WB 6.2 6.4 5.8 6 o8 6.5 .21 JB WB (c) JB 9.2 7.5 8.1 9.0 8.4 ,27 JB 8.0 8.6 9.0 7.8 8.5 .28 WB 5,6 6. 2 6.4 6.8 JB WB (d)d 6.2 .25 JB (e) JB 9.2 9.0 8.7 8.5 8.8 .20 JB (f) JB 8.4 8.8 9.1 7.6 8.5 .55 WB JB (a) JB 7.8 9.0 9.2 8.7 8.7 .51 WB JB (b) JB 9.1 7.2 8.7 8.1 8.5 .41 WB JB (c) WB 5.6 5.8 6.2 6.6 6.0 - WB 5.2 6. 0 5. 8 6.8 5.9 WB JB JB 8.5 9.0 9.2 7.9 ( d ) d .22 8.6 .29 WB (e) WB 6.5 6 04 5.5 6.2 6.1 .20 WB (f) WB 6. 4 6.7 6 .0 5.8 6. 2 .20 *See footnote to table 5. -24- Table 5. E f fe ct of d i f f e r e n t t r e a t m e n t s on the a s c o r b i c acid _______ content of f r ui ts~, 1947,_________________________________ ~ ^ '• Graft combination Treat­ ment s -a- Fruit A s c o r b i c acid, m g / 1 0 0 gm. analy. - of 4 i n d i v i d u a l pla nt s zed Sto ck - S c i o n 1 2 5 4 of f r u i t , M e a n + - S.E. WB * (a) WB S O .0 29.2 27.6 28.4 28,8 .52 JB WE (b) WB 28.4 28.8 28.0 27.6 ,26 JB WB (c) JB 26.0 2 4 . 4 28.0 26,6 25 . 5 .97 25.2 24 . 0 26.0 24.0 24 . 8 .49 JB WB JB * — JB 26.2 (d)Cl- ivB 27.2 28.0 o G .0 26.0 27.3 JB (e) JB 24.0 26 .0 25.6 27.2 25.2 .85 JB (f) JB 24.8 24 . 0 26, U 25.2 25 . 0 ,42 WE JB (a) JB 24.0 26.8 WB JB (b) JB 27.2 WB JB (c) WB WB JB __— tvB (d)C .85 22.8 25.2 24.7 .85 24 . 4 24. 0 26.0 25 . 4 .74 28.0 2 9 . 0 50 . 0 27.6 50.0 28 ,4 51.6 JB 24,0 26 .4 23.6 28.6 .54 26.0 27.6 1.20 26.8 25.2 .82 WB (e) WB 29.6 2 7 . 2 26.0 50 . 0 28.2 .92 WB (f) WB 30.0 28.0 28.7 .65 *See footnote to table 5. 27.2 29.6 -25- Tlie v a r i e t y W hi te B e a u t y lower i n total s o l ub le si gn if ic a nt l y h i g h e r was solids f o u n d to be and ti tr at ab le in a s c or b ic significantly acidity, acid conten t than Jo h n Baer, as is e v i d e n c e d f r o m the results u n d e r t r ea tm e n t s the u n g r a f t e d co ntrols. p o s i t i o n factors under t r e at me nt for (a) The v a l u e s fruits the s c i o n va ri et y use d. r o o t s t o c k was n e g l i g i bl e. which d e v e l o p e d u n d e r (b)), h a d more from the compa rab le other symbiont, ments. of the scion shoot, of stock lea' es ( t r e a t m e n t to that of the fruit (c), elaborated food materials from its solids, acids scion and in each, case un de r the result The shoots pro du ce d fruit on each, graft its own va ri et y. tion of fru it of plants influ enc e from the c o mp o s i t i o n the d e f o l i a t i o n of d id n o t com­ an d a sc or bi c acid for. t r e a t m e n t stock, and the in fl uen ce in an a p p r e c i a b l e (f). of change the I n fruit w h i c h c o u l d be a s s o c i a t e d w i t h e i t h e r st oc k or scion influe nce . ristic of that the fruits scions (f), that of the f r u i t of f r u i t fr o m c o nt ro l plants of fruit and st ock fruit f r o m trea tm en t c o n t a i n e d total In other words, composition, The obtained to the dev el op me n t The si mi la r scion var iety. scion leaves, shows the i n f l u e n c e or less which a p p a r e n t l y c l o s e l y with This (e) for these d i f f e r e n t o b t a i n e d fr o m the co m p a r e d and was not o f g r a f t e d plants , symbiont These with the results in t r e a t m e n t qu a l i ty (d), characte­ s h o w that the c h a n g e d b y any of the e x p e r i m e n t a l composi­ treat­ In st an ce s, on the qu al it y are quite in fruit trees, of the i n f l u e n c e s of s to ck and c o m p o s i t i o n of fruit bo rn e toy the scion, com m o n in h o r t i c u l t u r a l lit er atu re. The o b s e r v e d effects ha v e b e e n e x p l a i n e d e i t h e r on the basis of di re c t influence of s t o c k on the vigor the i n d i r e c t influence broug ht As d i s c u s s e d ea rlier, investigations were p e r f e c t l y o t h e r a n d the g r a f t e d p l an t s in vigor. It a p p e a r s one of the graft that symb io nt , fl ue nc e of the o t h e r tivi ty of t h ou gh t - At the in the g r e e n h o u s e . scope has b e e n p l a c e d m e n t i o n e d that the signs the of cha ng e fruits although developed under of the i n ­ ac­ to l i m i t controlled conditions considered necessary p r o b l e m by. a t t e m p t i n g to sta ge s by L y s e n k o the y o u n g e r to be also f ir st year's- work, the n u m b e r of v a r i e t i e s studies under more o f the desired (1947-48) c o n c l u s i o n of the It w a s at d i f f e r e n t w h i c h are grew w i t h o u t any s y m b i o n t m a i n t a i n e d the m e t a b o l i c desirable u s e d and p u r s u e plants compatible with each that v a r i e t y o Procedure crease the the v a r i e t i e s u s e d in this e x p e ri me nt , Experiment it was s c i o n or a b o u t toy the de gr e e o f c o m p a t i b i l i t y of stock a n d scion. i n these of the of g r o w t h . 011 the age of the plant, ch an ge d, Particular the the mo r e the to i n ­ graft the emphasis scion. characters He has of s u c c e s s f u l will toe -27- the ex p e r i m e n t . p la n t s On from which it is d e s i r e d or c h a r a c t e r m u s t be older; The w o r k was w i t h the o b j e c t the o t h e r hand, scions and comparatively st ab l e fo r d i f f e r e n t were sele ct ed , young ure was m o s t l y but due far as h u m i d i t y , ever, constant technique the older fall, if they are m i d d l e 1947, stocks. Two v a r i e t i e s w e r e ma d e w i t h o u t m u c h s u cc e s s temperature and vigilant and secure i n the g r e e n h o u s e , to to g r a f t v e r y at first. exercise proper and a e r a t i o n w e r e e f fo rt s conditions w h i c h are J o h n B a e r a n d W h i t e Beauty^ of attempts to i n a b i l i t y aged. i n t e r a c t i o n of v e r y y o u n g characters, number a seedlings, it is b e s t studying c l a i m e d that the to o b t a i n a c e r t a i n p r o p e r t y started during of h e has d i d he l p favorable The fail­ controls concerned. to d e v e l o p as How­ the for e s t a b l i s h i n g the graft u n i o n s . During January, of J o h n B a e r twenty successful and W h i t e B e a u t y w e r e m ad e, l u s t r a t e d in f i g u r e s i s t e d of s i x - w e e k were 1948, r e m o v e d just s i s t e d of small II. The at the seedlings using stock plants old tomato plants time two w i t h f l o w e r buds, The cotyledons g r af t s the m e t h o d il ­ in each, case of g r a f t i n g . with reciprocal sci on s an d two con­ which con­ small leav es. The scio ns the d e v e l o p i n g in a l l scions these gr af t s grew, presented peculiar but the l e a v e s symp to ms . of Th e y w e r e -28- thick, curled and deformed. malities were not rej ec te d. met the favor same type attempt Avakian and Jastreb ful reciprocal made, using year. three u s e d as stocxs The weetcs o l d p l an ts . spaced In addition each variety, of These the during and g r a f t e d plants, age as for the p u r p o s e t r a n s p l a n t e d gr a f t s as the treatments control tr e l l i s e s . These greenhouse for the s i x weeks o l d and o b t a i n e d from were the first set w e e k of the pla nt s in the with fruit were trellises were of stocks, the fruit fr o m u n g r a f t e d plants. showed st ar te d. pl an t s w e r e five plants of c o m p a r i n g So o n a f t e r tablishment, the p r e v i o u s those u s e d for symbionts the success­ apart. the sa m e transplanted year, of eighty grafted plants 5 fe e t a p a r t feet to in and Wh it e B e a u t y were were f r o m the g r a f t we l l 1948, sci on s we r e in five rows rows w e r e given up (1941). in h e i g h t . The seedlings the e x p e r i m e n t of J o h n B a e r June. abnor­ were the p r e v i o u s the t e c h n i q u e f o l l o w e d d u r i n g cm. were again grafts greenhouse we r e once part of Ap ri l , in the rows pl a n t s and h a d to be last Th e p l a n t s 25 to 5U and the followed during to r e p e a t of these to g r a f t y o u n g e r of r e s u l t s of the m e t h o d D u r i n g the causes a c c o u n t e d for, Another with a view The signs The t r a i n e d to especially of grafted two es ­ as sh o o t s on set up' in the purpos.e of p r o v i d i n g u n i f o r m l i g h t -29- condi ti on s of close v.hi oh c ou ld not be p r o c u r e d o t h e r w i s e plan ti ng . sh oo t f r o m the grow. The following (a) was In types from treatments the o ne stock we r e a l l o w e d a d o p t e d c o n s i s t e d of to the of training: In the case of u n g r a f t e d d e f o l i a t e d and the under the case of g r a f t e d plants, scion and one different because the i n f l u e n c e fruit c o nt ro l pl ant s, allowed o f the lea ve s of to set o ne sho ot and develop the o t h e r shoot, which was d e f l o r a t e d 0 (b) were The shoot a l l o w e d to stock lea ve s (c) The the i n f l u e n c e and d e v e l o p shoot the o f the fruit, which but (e) on The was the leaves All v e g e t a t i v e shoot was scion shoots. developed under of the stock. set a n d d e v e l o p were treatments to set g r o w t h was of fruit u n d e r n ot a l l o w e d to s e t kept intact, defoliated and allowed the i n f l u e n c e allowed of c o n s i s t e d in d e f o l i a t i o n of the each of the a b o v e of f l o w e r s s t o c k and s c i o n w h i c h was s t o c k was d e v e l o p fr ui t u n d e r In on bo th and a l l o w i n g it to the i n f l u e n c e flowers influence d e f o l i a t e d an d the fruit f ru it on the s c i o n treatment the the s t o c k fruit, of leaves a n d fruits This d e f ol ia te d, set f r u i t a n d m a t u r e u n d e r s c i o n s h o o t was In o t h e r words, sto ck s c io n was but in the a b s e n c e of a l l o w e d to set (d) f r o m the to set a n d s c io n l e a v e s only fruit on the and first c l u s t e r the d e f o l i a t e d s u p r e s s e d a f t e r the fruits. shoot. fruit h a d set. The number* o f pla nt s u n d e r d i f f e r e n t treatments are p r e s e n t e d i n table 6. T a b l e 6. N u m b e r of u n g r a f t e d and g r a f t e d pl a nt s u n d e r _________________ d i f f e r e n t t r e a t m e n t s emp l o y e d d u r i n g 1946. Graft combination No, of N u m b e r of g r a f t sy m b i on ts ungrafted u n d e r d i f f e r e n t tr ea t m e n t s _______________________________ p l a n t s _____________________________________ S to c k - c»cion (a) Jo h n B a e r White White Jo hn Baer Beauty Beauty 5 t re at me n t s characters under (e) 5 10 5 5 10______ 5 10 40 and D i s c u s s i o n - The under different (d) 20 10 Results (c) _5 _ ______ 20 T ot al the (b) fruits 20 of graft d e v e l o p e d no r m a ll y. study were 10 s y m b io nt s No e x h i b i t e d b y fruit ch an ge s in from g r a f ­ ted plants. The fru it s u n d e r st ock l e a v e s of s c i o n used. which in the fru it s was The frui ts (c), ex ac t l y w i t h of its studied, were fr o m the the we re n o r m a l the ( b ) , wh ere ab se nc e aeveloped under treatment treatment the influence of fruits on the development typical of the scion d e f o l i a t e d sho ot s influence of s t o c k frui ts a n d c o m p a r e d in s c i o n v a r i e t y used. of the va ri et y scion, and leaves, f o r m a n d color -61- The influence treataients influence (d) of as rived and scion (e) . (d) stock but and under fr ui t u n d e r on s t o c k The scion leaves in t r e a t m e n t well of fruit fruit in t h e the treatment absence The elaborated food materials m o s t l y did n o t present-any These trolled results conditions treatments, change in the of characters and observation the findings of scion the a s c o r b i c ence in varieties are k n o w n acid content ascorbic acid of their content the entirely under genetic tent o f of a parti c u l a r vi di n g fruits roots To s t u d y these questions, grafts greatly fruits. Various given secure to control? the variety in the an e x p e r i m e n t were Is expression made ascorbic of of conditions for of a was differ­ acid by content two of pro­ in varieties their later, investigating con­ variety? designed of in character another treatments,- to be d i s c u s s e d the this changed plants acid Synthesis soluble ascorbic be con­ year. considerably fruits. Gan study. experimental first differ or p h o t o synthesizing p a r t s which r e c i p r o c a l differing to leaves carefully Stock-Scion R e l a tionship with R e f e r e n e e to o f A s c o r b i c a r i d a n d i t s d e l a t i o n to T o t a l Solids and Acidity Tomato as -which d e ­ under of the fruit leaves fruit under the scion the in under scion from obtained confirmed of stock which were strict studied, developed influence (e) . \ 7, indicate a a n d 9. 8 John Baer and ' Y e a g e r 1 varieties res­ Treatment: (a) One s t e m v i t h fruits; u n g r a f t e d p l a n t (in gr e e n h o u se ) (b) Tvvo sh oo ts , one f r o m s c i o n and one f r o m stock; s c i o n d e ­ f o l i a t e d a n d its fr u i ts d e v e l o p e d u n d e r the i n f l u e n c e of s t o c k l e a v e s ( ot h e r v a r i e t y ) . (c) One s h o ot f r o m s c i o n w i t h fruits; st oc k p r o v i d e d roots only. (d) Two sh oot s, one f r o m s c i o n and one f r o m stock; s c i o n d e ­ folia te d; its f r u i t s d e r i v e d f o o d m a t e r i a l s f r o m st ock l e av es (same va ri et y) . (e) One s t e m w i t h fruits ; u n g r a f t e d p l a n t (in f i e l d ) „ Note: (a), (b), (c), a n d an d (e) in field. (d) treatments p e r f o r m e d in g r e e n h o u s e -4 2- Table 8. To t al s o l u b l e solids content of ripe fruits of to m a t o e s f r o m different treatments of two graft c o m b i n a t i o n s , 1948._________________________________ Trea truent# Graft combination Stock solids plants Me an of 1 2 5 4 5 5.0 5.6 4.8 5.2 5.4 5.2 .14 .10 Scion JB (a) Per cent t ot al soluble fruits of 5 i n d i v i d u a l t S o -t-J F • O (b) Y JB 5.0 5.4 5.2 5,0 CO • 5.1 (o) Y JB 5 .2 I. 8 5.4 5.6 5.0 5.2 .14 (d) JB JB 5.2 5.0 5.0 4.8 5.0 5.0 .02 (e) JB 7.8 6 .8 6.0 6.4 7.0 6.8 .50 (a) Y 4.0 4. 4 4.2 4.0 4.4 4.2 .09 (b) JB Y 4.2 3.6 4.4 4.6 3.8 4.1 .19 (c) JB Y 4.4 5.6 3.6 4.0 4.5 4.0 .19 (d) Y Y 4.4 3.8 4.0 4.0 3.8 4.0 .11 6.0 6.0 5.6 4.4 5 .0 5.4 .51 Y (e) wS e e footnote t a b l e 7. -45- Table 9. Titratable acid content of ripe fruits of tomatoes from different treatments of two __________ graft combinations, 1946.___________________ O Graft Treat­ combination me nt* - Scion Stock JB (a) A c i d i t y in terms of ml. of 0.1 i'l Na O H u s e d to titrat e a 20 ml. a l i q u o t 2 5 4 10 . 4 9.2 8.7 9.0 1 5 Mean ±3.b . 10 . 0 9.5 .52 0>) Y JB 8.7 9.5 10.0 9.5 10.5 9.6 .28 (c) Y JB 9.7 10.4 9.1 10.2 8.7 9.6 .52 (d) JB JB 8.8 10.7 10 . 2 10.0 9,6 9.9 •5 2 1 1 . 0 11.5 9.0 11.5 10. 5 10.7 .45 7.7 7.4 8.0 7.1 8.2 7.7 .20 (e) JB (a) Y >(*>) JB Y 7.5 8.0 8.6 7.4 9.0 8.1 .51 (c) JB Y 7.5 8.1 6.1 8.2 8.0 7.6 .59 (d) Y Y 7.5 7.1 8.8 8.6 8.5 8.1 .54 5.9 6.6 7.5 o.5 7.0 6.7 .27 (e) tf-See f o o t n o t e Y t a b l e 7. -44- S t o c k - S c i o n R e l a t i o n s h i p in .Connection W i t h S t u d i e s on R e c i p r o c a l G r a f ts of N o r m a l Plants an d PTanTs M o d i f i e d T h r o u g h the a c t i o n of 2 , 4 - d i c h l o r o p h e h o x y a c e t i c ~ . a c i d The malformations the ir similarity somal disturbances since their compounds to virus m a n i f e s t a t i o n s (Stei n 1959), observation. act a n d p r o d u c e cal ,d i s t u r b a n c e s The p r o d u c e d b y p h e n o x y com po un ds , ha v e (1948). Stein (1959) and i n d u c e d c h r o m o ­ b e e n of i n t e r e s t these physiological and m o r p h o l o g i ­ workers t h r o u g h graft u n io ns such as V a s u d e v a o b s e r v e d th at g r a f t i n g of a .tomato mu ta n t , character variations w h i c h r e s e m b l e d 2 , 4 - D i n d u c e d c h a n g e s ) , onto tomato plant lo g i c a l plants, c a u s e d its characters The o b j e c t of the and c h l o r o p h y l l and S a m Raj wit h d w a r f mal deficiency elongation. by the u s e of 2,4-D, would a nor­ o th er m o r p h o ­ remained unchanged. i n v e s t i g a t i o n was i n which, t y p i c a l m o r p h o l o g i c a l (radi um i n d u c e d However, of the g r a f t e d m u t a n t present ever is y e t u n k n o w n . t r a n s m i s s i o n of c e r t a i n vi r u s es shov.n by v a r i o u s with The m e c h a n i s m b y w h i c h the p h e n o x y in plants has b e e n . to s t ud y h o w the chang es h a d be e n i n d u c e d react wh e n grafted with normal plants„ procedure Plants 40 cm. of the J o h n B a e r in height, were variety, t r e a t e d by five w e e k s immersing their o l d a n d 50 to third basal leaves in a 1000 p. p . m. lorophenoxyacetate aqueous sodiu m id,4-dich- d u r i n g the first w e e x of N o v em be r, The n e c e s s a r y p r e c a u t i o n s , parts o f solution of the p l a n t s , we r e to p r e v e n t 1947» c o n t a m i n a t i o n of o t h e r t a k e n by c o v e r i n g the t r e a t e d leaves w i t h cellophane b a g s c After displaying the u s u a l n a s t i c eff ect s, developed m a l f o m a t i o n s after th e tr eatment, reciprocal of splice grafting grafts on when their t e r m i n a l grafting, i l l u s t r a t e d in fi gu r e o f each c o m b i n a t i o n w e r e plants e a c h of the not s e v e n to t e n days a l l o w e d to d e v e l o p a l l o w e d to g r o w weeks after were noticed successful In addi ti on , were twelve allowed found estab­ -axillary sho ot s were the s t o c k d u r i n g However, Before this p e r i o d and s u b s e q u e n t l y the p l a n t s and D i s c u s s i o n the u n i o n s h a d d e v e lo pe d, ted g r o w i n g a p p r e c i a b l y . fluences ob tained. a f t e r gr a f t in g, The m e t h o d without being disturbed. Results Th ree Twelve The g r a f t u n i o n s were from for two we eks a f t e r w a r d s . were v! ;ell e s t a b l i s h e d , t r e a t e d a n d u n t r e a t e d groups, their o w n roots. weeks I, was u s e d . w e r e re moved. grafts l is h e d Three w i t h u n t r e a t e d plants. the t r e a t e d le a v es to g r o w o n parts. th os e e f fe ct s w e r e were made all pl an t s At that time on some n o r m a l signs scions sci on s star­ of f o r m a t i v e g r a f t e d on in­ stocks -46- in- w h i c h m o r p h o l o g i c a l w i t h 2,4-D. The changes h a d b e e n i n d u c e d b y symptoms after g r a f t i n g were e v i d e n t on the pla nt s recorded. Quantitatively, si derable differences the gr af t combination where normal tr ea te d stocks. noticed. in the r e s p o n s e of Ty p i c a l plants three m o n t h s there w e r e individual sc i o n s we r e Q u a l i t a t i v e l y , .however, treatment p l an ts of g r a f t e d on a definite of ea c h gr ou p a r e con­ t r e n d was shown in fi gu re s III and l'V. P la nt 1. a n u n t r e a t e d plant. No p e c u l i a r i t i e s were evi­ dent . Plant 2. of leaves a t r e a t e d plant. a n d even fruit s. p l a n t o. o r i g i n a l l y was a normal those few lea ve s gans were Plant stock was grafted with a scion which the a p i c a l p o r t i o n of a t r e a t e d plant. and a r r o w in di c a t e m o v e d from It s h o w e d e x t r e m e m o d i f i c a t i o n s the g r a f t u n i o n , plant, tag 'with the e x c e p t i o n of w h i c h were m o d i f i e d b e f o r e the o r i g i n a l The the s c i o n was no m o r p h o l o g i c a l l y re­ changed or­ r e c or de d. 4. a treated o ri gi na ll y was the foliage of the s to cx was g r a f t e d with a s c i o n w h i c h api.cai p o r t i o n of a n o r m a l plan t. scion exhibited morphological which i n d i c a t e d that the f o r m a t i v e m i t t e d t h r o u g h the g r a f t union. The abnormalities s t i m u l us h a d b e e n trans­ Pig. Ill P l a nt 1. U n g r a f t e d , untreated. Plant 2. U ng ra f t e d , treated. Figure III Pig. IV. Plant d. S ci on f r o m a t r e a t e d p l a n t onto a n o r m a l stock. Pl an t 4 0 S c i o n f r o m a n o r m a l a tr e a t e d stock. arrow indicates plant grafted g r a f t e d onto the g r a f t - u n i o n . Figure IV -47- P r o m the experimental greater quantities portion and fr o m there It seems treated the could scion, st oc k (Plant 4), came u n d e r ungrafted plant like the t i c e d th a t cut s t um ps the phenoxyacetic 50 da y s of the changes chi ci ne . chemical could have They further shoots tips wer e a p p l i e d to the in­ induced by col­ s u g g e s t e d that i n f l u e n c e we r e o l d stem if the substance must findings i n d u c e d since a normal of d i r e c t scio ns u s e d f o r g r a f t i n g indicate co n t i n u e of to m o v e the tip o f the t h a t no g e n e t i c the m o d i f i e d s c i o n w h e n g r a f t e d on s t o c k d e v e l o p e d into to be the o c c u r r e d su c h as w e r e The p r e s e n t seems it was gene­ a n d th e n u p w a r d to influence with modified or w h e t h e r in the a normal no­ stump downward were the lower (1942) still u n d e r the c ha nge s t r e a t e d but T h e y m e n t i o n e d that chemical nature shoot. the of treated with p-chloro- a direct new influence 'which s u b s e q u e n t l y d e v e ­ of tomatoes the g r o w i n g constant and H i t c h c o c k a f t e r treatinent. s to ck it h a d u n i t e d w i t h the parts of acid produced many new n ot k n o w n w h e t h e r tic Zimmerman that scion. st im ulu s m o v e d from to the y o u n g g r o w i n g parts leaves . fluence The the n o r m a l the t er mi na l (Plant 2). loped modified le av e s , after appear s t o r e d in the influence the n o r m a l just it w o u l d s t i m u l i we r e that stimulus portion of the results, top' (Pla nt 5). c h e m i c a l na tu r e . (Pla nt 5) e i t h e r d i d n ot The The m o d i f i e d carry any -46- f o r ma ti v e itself s t i m u lu s or it was in the f o r m of m o d i f i c a t i o n s No p o s i t i v e stimulus, s ta tem en t tion, after to express in new growth. can be m a d e as m e n t i o n e d by Z i m m e r m a n only a few ca ses have cal was too i n s i g n i f i c a n t abo ut the f o r m of the an d H i t c h c o c k (1948), b e e n r e c o r d e d in v'hich the a p p l i e d chemi­ d e t e c t e d or i d e n t i f i e d b e y o n d the r e g i o n of a p p l i c a ­ The p r e s e n c e of 2 , 4 - D in p l a n t s has n o t b e e n shown its a b s o r p t i o n . However, n u m e r o u s wo rk e r s h a v e r e p o r t e d on a b s o r p t i o n a n d t r a n s l o c a t i o n of 2 , 4 - D d e r i v i n g their con ­ clusions from indirect and Brown (1946) have evidences s u g g e s t e d that associated with that of o r g a n i c the fate of the su b s t a n ce tion tuid p a r t i c u l a r l y phological changes during the take place. foods. tained for three week s, un ti l the or a d e r i v a t i v e attempts the g r a f t u n i o n and of the scion. Ho wever, c o nc lu s i v e l y , the c a u s a t i v e it is a a g e n t was 2,4-D o r even a r e a c t i o n pr od uc t, w e r e made under which 2,4-D which moved t r a n s l o c a t e d and re ­ c o n c l u s i o n is p e r m i s s i b l e of the leaves of s p e c u l a t i o n w h e t h e r and t r a n s l o c a ­ the a s s u m p t i o n that 2 , 4 - D s u c h an a s s u m p t i o n ca n be p r o v e d matter o f 2,4-D is N o t h i n g is k n o w n a bo ut a b s or be d, that it was 2 , 4 - D that m o v e d t h r o u g h caus ed the m o d i f i c a t i o n Mitchell subsequent period when m o r ­ Under e x p e r i m e n t was such the t r a n s p o r t af te r its a b s o r p t i o n in the p r e s e n t as of g r o w t h re sp o n s e s . to test the v a l i d i t y as s u c h c o u l d be the t h r o u g h the g r a f t union s. of the form o f the Direct assumption stimulus ap pr oa che s, by -49- tryi ng to i s o l a t e whether 2,4-D and detect 2, 4 - D , were m a d e c o u l d be abso rb ed , to investigate t r a n s l o c a t e d and r e t a i n e d as suc h for s u f f i c i e n t l y lon g per iod s. The result s fi ndings do no t lie of the st udi es w i t h i n the he r e and w i l l be p r e s e n t e d How ev er , it m a y be fi n d de te c t a b l e its the present in a sep ar at e p a p e r reported e l s e wh er e. s t a t e d that it h a s not b e e n p o s s i b l e to quantities a p p l i c a t i o n to scope of those of 2,4-D the le a v e s investigations0 in tomato p l a n t s in c o n c e n t r a t i o n s after as u s e d in -50- GENERAL DISCUSSION The studies on s t o c k - s c i o n int er ac ti on , w h i c h have Le e n r e p o r t e d in the st art ed in order to analyze pr ev io us (1946) ted. results The after this binations of g r a f t e d pla nt s tried. The fruits and t it ra ta bl e e x p e r im en ta l sym bio nt influence of were of ability of the the fr u it s were not disturbed by which to d e v e l o p u n d e r However, e n o u g h to dis pr ov e L y s e n k o has and s ci on occ ur s H e has to f o r c e by the vai'ietjr, the properties He h a s the "nutrient" the claimed that o n l y und er s t a t e d that the s c i o n to ap­ fre­ the "assi mil ate " ma t e r i a l s m a n u f a c t u r e d o f w h i c h are further added the these re su lt s v e g e t a t i v e h y b r i d s would, d e p e n d u p o n of the scion. fruits sy mb ion t. c o n di ti on s. that c o n s i s t e d in d e f o l i a t i n g conclusive experimenter com­ indicated as m a n y as possible to the any graft total so lu b l e the i n t e r a c t i o n b e t w e e n s t o c k que ncy of o b t a i n i n g in f r u i t of a s c o r b i c .acid, vegetative h y b r i d i z a t i o n co nc e p t . propriate b i o l o g i c a l ch ang es fin dings the o t h e r g r a f t to be No with r e g a r d a n d a l l o w i n g its not c o n s i d e r e d inia- These treatments, one graft w o r k h a d be e n were n o t i n f l u e n c e d acidity. the m e t a b o l i c p r o c e s s e s the con ­ w i l s o n and we re n o t i c e d u n d e r to their c o m p o s i t i o n in ter ms solids were the f i r s t y e a r s work d u r i n g 1947 c o n ­ fi rm e d W i l s o n and W i t h n e r 1s f i n d i n g s 0 characters of the v e g e t a t i v e h y b r i d i z a t i o n paper appeared of results se ctions, cept since no other wo rk h a d b e e n reported. Yiiithner's the to be transmitted that in m a k i n g grafts, the -51- s ci on s h o u l d be "age", as y o u n g as po s s i b l e m e n t i o n e d b y H u d s o n a nd hi ch e n s as t e r m i n o l o g y of n u t r i e n t s Nu tri en ts , ment al to him, m a y c o nd it io ns , su bs ta n ce s n e c t i o n wi th he me n t i o n s all the b a s i n g his four of Pla nts". or ga n is ms he uses types vegetative hybridization f ol lo we rs , as ages on hi s Considering this, Attempts made d u r i n g 1948 g ra ft in g) , and two to scion seems that w h e n and sc io n he m a y be of "p has ic D e v e l o p m e n t c o mp le te ana lys is o f the difficult. Lysenko t r a n s l a t i o n of very y o u n g bef or e the ir to knowclaims to graft v e r y y o u n g scions buds though successful, techni­ It w o u l d be d e s i r a b l e small l e a v e s w h i c h the f l o w e r teraction between sto cK experimental procedures adequately. fr o m or or gan ic to s u p p l y d e t a i l e d i n f o r m a t i o n on the can be t e s t e d m o r e old, of It r e v e a l e d by the E n g l i s h of experimentation. cotyledons f r o m s to cx environ­ t e r m a s s i m i l a t i o n in c o n ­ concept becomes cal as pec ts two the the fail their such as gam et es , co n c e pt their work, more a b o u t su bs t a n c e s , of "n ut ri en ts ". comparative terminology Lysemto's and a s s i m i l a t i o n is v e r y vague. s u p p o s e d to pass Similarly, stock of medium, (1946), be e i t h e r i n o r g a n i c whole s u c h as are in gr afting. and his and the were onto o l d sto ck s r e m o v e d at the d i d not h e l p scions in the (six weeks time of st udy a n d o l d stocks. with of i n ­ The scions soon after t h e i r e s t a b l i s h m e n t , p r e s e n t e d p e c u l i a r s y m p t o m s gr o w t h r e s e m b l i n g virus m a n i f e s t a t i o n s , be r e jected. and the p l a n t s in h a d to xi r e p e t i t i o n of the e x p e r i m e n t which c o n s i s t e d in g r a f t i n g old toma to pl an ts results the p r e v i o u s year. of of these re su lt s, of the type that stages of the p r e s e n t finite conclusion can be studies c o n f i r m e d the its cruci al experiments. rep e t i t i o n s of the st oc k and scion, is u n l i k e l y w h e n pla nt s are too c l a i m that sc an t y the been d i s p r o v e d . d i s c o v e r y has three-week s t a t e d on the b a s i s of g r o w t h m e n t i o n e d . ha v e However, to the re­ ju st if y a d e ­ findings The are of the pr oo f of a alw ay s b e e n in the r e p r o d u c t i o n of It is experiments support to L y s e n k o * s obtained, fr om old plants, It d u r i n g 1948, an i n t e r a c t i o n b e t w e e n and any Russian scientists sci en t i f i c six-week c l a i m e d by Ly se n k o , g r a f t e d at the sults onto sc i o n s of 1947, only b y ca r e f u l w h i c h are t h e or ie s, that the a d di t i o n a l s u p p o s e d to le n d c o n t r o v e r s y wi l l be resolved. C o n s i d e r i n g L y s e n k o ’s t h e o r y Plants" w h i c h is complete rejection, the e x p e r i m e n t s on the b as is plants. cause This of the di f f i c u l t b e ca us e changes; the to su b j e c t of 11Ph a s i c d e v e l o p m e n t by of p h y s i o l o g i c a l s e l e c t i n g the sele ct aspec ts pl an t s to r e p e a t stock or developmental approach w o u l d involve technical of some d i s a g r e e m e n t but not it w o u l d s e e m d e s i r a b l e on g r a f t i n g new of of d i f f e r e n t ages of the work b e ­ It is r a t h e r d e v e l o p m e n t a l ages these m a y n o t be a s s o c i a t e d with a n y m o r p h o l o g i c a l and p r o f o u n d i n t e r n a l developmental of an d s c i o n considerable of the problem. some ch a n g e s m a y be -55- re vs ale d only u n d e r d e f i n i t e environmental conditions,, The p r o b l e m is f u r t h e r c o m p l i c a t e d by the fa c t methods ex ist to a s c e r t a i n tive cha nges. It would under s t r i c t l y w hi ch are an e s s e nt i al The studies extended by prerequisite it on the find out would b e no e v i d e n c e from one results corbic of content fruits of to to ass um e phenomena. that a tomato differ­ as i n d i c a t e d the basis s ub st an ce s of some role of a n s ci on or in g r o w t h W i l s o n and v.ithner (1946) thiamin, found r i b o f l a v i n and n i a c i n the o t h e r in g r a f t e d tom at o plants. the p r e s e n t i n v e s t i g a t i o n s a c i d gave no On strong variety develop un d e r be vdiich h a v e a l r e a d y b e e n re ­ substances may play to and s ci on m a y t r a n s m i t t e d from s t o c k to of t r a n s f e r o f sy mb i o n t con di ti on s the p h y s i o l o g i c a l s u c h as vitamins. logical c a n be These and d e v e l o p m e n t a l ex­ d e t e r m i n a t i o n of and u n g r a f t e d pl a nt s co ns tituents, ve r s a . for the i n t e r a c t i o n of stock a t t e m p t i n g to a l k a l o i d a l n ature vice these of plants,, the f i n d i n g s of som e wo r k e r s , viewed, qualita­ c o n t r o l l e d en vi ro n m e n t a l ences b e t w e e n the g r a f t e d by c e r t a i n these in ternal b e n e c e s s a r y to c a rr y out periments p h y s i o l o g i c a l age some o f that no e a s y evidence on the that could be the sy nthesis ascorbic The of as­ acid c h an ge d by a l l o w i n g its the i n f l u e n c e of the leav es of an o t h e r variety.. Although need for Lysenko's hypotheses thorough suggested. analysis are open to argument, a of his g r a f t i n g e x p e r i m e n t s has It is c o n s i d e r e d that a n y facts revealed been by fu rt he r -54- r e p e t i t i o n of his ex per iments, the r a p i d l y p r o g r e s s i n g fie ld s and p h y s i o l o g i c a l B'yfe (1947) c o u l d be of d e v e l o p m e n t a l p h y s i o l o g y genetics. ha s s t a t e d that th e n e w p r o p o s e d by L y s e n k o may be must b e a d e q u a t e l y yyhyte and c a r e f u l l y (1946) the R u s s i a n w o r x e r s th eo r i e s b ut teste d, that his before seexaed to a p p r e c i a t e in t he ir the re s u l ts on g r a f t i n g exchange in r e la ti on to h o r m o n a l (Hamner a n d B o n n e r 19 58 ), possibilities of the ex pe r i m e n t s a ve r d i ct findings grafting experiments. p r o p o s e d a c o n n e c t i o n b e t w e e n these ing of g e n e t i c s en t i r e l y w r o n g and m a y be d i s r e ­ g a r d e d as a c o n t r i b u t i o n to bi ol o g y , passed. i n c o r p o r a t e d into these of He has and the r e s e a r c h to induce and e v e n s p e c u l a t e s a p p l i c a t i o n of is resul ts flower­ about the to crop producti on , we n t scion, (19o8), using different varieties gr ow t h of the fect, He in i n v e s t i g a t i o n s of the ef f e c t of g a r d e n pea, s ci on was n o t m e r e l y a c o n s i d e r e d that the in the sible si m p l e n u t r i t i o n a l presence r o o t s t o c k w h i ch he n a m e d " c a l i n e s 11. that in tomatoes such su bs t a n c e s end may in f l u e n c e in m o r p h o l o g i c a l over of such m o r p h o l o g i c a l l y d o u btf ul . the ef­ ch ara cte rs in of g r ow t h factors It is quite p o s ­ of h o r m o n a l n a t u r e m ay be pr es e n t su lt in g g e n e r a t i o n is f o u n d that e x p r e s s i o n of ma n y the s c i o n a p p a r e n t l y d e p e n d e d on the of s t o c k on the changes. s t o c k - s c i o n reactions, The c h an ge d However, re­ p o s s i b i l i t y of a c a r r y ­ characters thu.-re is into evidence the ne x t that some -55- of the ef fec ts i n d u c e d by s y n t h et ic g r o w t h r e g u l a t o r s c a r r i e d t h r o u g h s e e d into and W i t t w e r 1949). The the f o l l o w i n g g e ne ra ti on s q u e s t i o n of how this are (Clark influence is c a r r i e d t h r o u g h is yet not decided. Strasburger as quoted by Batchelor found protoplasmic connections cells a n d those of the of the it is thus stock possible It does or o r g a n i z e d p r o t o p l a s m i c st oc k or vi c e versa, enzymes m a y be re actions. but The r e s u l t s formative acetic acid, not so of the the !lh o r m o n e - g e n e " Ho we ve r, Schmuck "hormone-gene" inter ac ti on , is an ex i s t e n c e o f the g en es genes are n o t in er t characteristic on the transmission s u gg es t that o t h e r s i m i l a r the gr af t union. in p h y s i o l o g i c a l ge ne t i c s about ge n e S o m e of the re su lt s cl a i m e d be e a s i l y e x p l a i n e d on the b a s i s p r o p o s e d by G o l d s c h m i d t has m e n t i o n e d that inadequate especially and induced by 2,4-dichlorophenoxy- hypothesis (1945) such as h o r m o n e s studies wide range. by R u s s i a n w o rk er s m a y fr o m s c i o n to and pr o d u ce s t im ul i c a n be t r a n s l o c a t e d ac r o s s a c t i o n cover a very s c i o n in c e r t a i n plants; w o u l d pass t h r o u g h gr af t un i o n s , The m o d e r n c o n c e p t s b e t w e e n the se em l i k e l y that pla sti ds substances stimulus (1948) is m i g r a t i o n of i d i o p l a s m bod ie s e x p e c t e d to do of the (plasmodesms) that there and plasm p a r t i c l e s . and W e b b e r the e x p l a n a t i o n of in vi e w of his opinion is ve r y do u b t f u l . protein molecules The of (1938). t h e o r y of the stock-scion that the act ua l c o n c e p t i o n that but p l a y an ac ti v e role -56- in the p h y s i o l o g y of cells, stances of h o r m o n a l p r o b a b l y by r e l ea si ng or e nz ym i c natur e, pr o c e s s e s of d i f f e r e n t i a t i o n biologist assume that the 1940) h as put ti on o f re g i o n a l essential by the e m i s s i o n of "hormones" react s pe tence. according tion in the tissues; the de ta il s of the govern at s p ec if ic it w ou ld be n a t u r a l a n d cha ng e of cells its identical fer divergences plasm has suggested D a c h of these of the o r g a n i s m or c o m ­ local d i f f e r e n c e s are in determina­ r e s p o n s i bl e for In the li gh t of s u c h 'c o n ­ to ass um e that c o u l d di ff us e throughout s ub st an ce s f r o m one the s o m a t i c the body; in d i f f e r e n t i a t i o n . that c o n c e r n e d in these the d i f f e r e n c e s abo ut w hi ch cell to on to g e n e s i s . genetically (1948) is b r o u g h t of maturity "hormones" It is g e n e r a l l y b e l i e v e d that ex tr em e the d e t e r m i n a ­ the bacic of d i f f e r e n t i a l individuation. the d e s t i n i e s another stage to this n o t i o n , of m a t u r a t i o n lie cep ti on s " h or mo nes ", appropriate the Goldschmidt that in tur n with that part r e a c h e d the rates co nt inu it y. in d e v e l o p m e n t successive w h i c h has the c o n t i n u i t y w i t h i n the f o r w a r d the view c h a ra c t e r s w h i c h control and m o r p h o g e n e s i s , m a k e s l i v i n g o r g a n i s m is a ch e m i c a l (1958, sub­ "ther e m u s t be ce l l u l a r cells are and jret they s u f ­ Wigglesworth ele men ts in specializations, the c y t o ­ 'that in cell h e r e d i t y that a r i s e w i t h i n a mult i- c e l l u l a r o r g a n i s m are cytoplasmic'." Darlington .(194a-) and 'Aright (1945) cytoplasmic d e v e l o p e d the cons t i t u e n t s , w h i c h t h e 3*- n a m e d D a r l i n g t o n has that n u c l e a r dington linked up heredity genes (1948) on importance nature yet cy t o p l a s m i c factors in m o r p h o g e n e s i s is of in the The fr o m s t o c k to Haldane Later, s c i o n or vice v e r s a has Lspinasse in 1944, (1944) (1941) Haldane in regarded that graft u n i o n was tion are thus to bu s h hab it , the ca u s a t i v e rather could not agent remar ks causes a re­ l e n d some wo rkers. sup­ Darlington transmitted through t h a n a virus, since the be c o n s i d e r e d p a t h o l o g i c a l . s t a t i n g that studies the explanation. might o b t a i n e d by the S o v i e t technical s t o c k or s c i o n s u g g e s t e d that D a r l i n g t o n ' s a r t i c l e by exist b e t w e e n the r e su lt s m i g h t o p p o s i t e m e m b e r of supported Haldane's a plasmagene in r o s e s c o n c l u d e d his fr o m the roses w h i c h in some cases the r e s u l t s bushy habit If a l r e a d y b e e n discu ss ed . s u g g e s t e d that L y s e n k o ' s v e r s i o n of c l i m b i n g h a b i t to c y top la sm. sc ie n t i s t s b e c o m e s t h r o u g h the i n v a s i o n of e i t h e r on b u d - g r a f t i n g port plants a c c e p t e d t h e n the e x p l a n a t i o n of by a n o n - p a t h o l o g i c a l virus graft. of some p o s s i b i l i t y of m i g r a t i o n of p l a s m a p a r t i c l e s (1940) be e x p l i c a b l e VVad- a self-reproducing s t o c k - s c i o n i n t e r a c t i o n c l a i m e d by S o v i e t evident. by sh owi ng and interact. sta ted that c e r t a i n c h a r a c t e r s self-reproducing bodies of "plasaiagenes1’. and d e v e l o p m e n t and p l a s m a g e n e s b o t h has are d e p e n d e n t the idea of s e l f - r e p r o d u c t i o n in of h e r e d i t y , and a r b i t r a r y "The frontiers development He that and i n f e c ­ and n e w p o s s i b i l i t i e s of an al y s i s and e x p e r i m e n t will the pa s s w o r d s to take us arise w h e n we have lea rn t across". T he se considerations, that no real d i s t i n c t i o n can be d r a w n b e t w e e n c y t o p l as mi c inh er i t a n c e results of an d virus in fection, make the e l u c i d a t i o n of the R u s s i a n wor k on v e g e t a t i v e h y b r i d i z a t i o n difficult. It is also difficult h y b r i d i z a t i o n a n d the of a g r a f t b y logic al to d i s t i n g u i s h b e t w e e n v e g e t a t i v e i n d u c t i o n of m u t a t i o n in one m e m b e r the other. Kostoff (1929) observed physio­ i n t e r a c t i o n b e t w e e n s to ck and s c i o n of c e r t a i n so l a na ce ou s species a c c o m p a n i e d by m o r p h o l o g i c a l He e x p l a i n e d the r e s u l t s by p o s t u l a t i n g The p o s s i b i l i t i e s Sturtevant c o n s i d e r e d that, p o n s i b l e for possibility the that react w i t h the (194a) by antibodies an d E m e r s o n "if a p a r t i c u l a r gene is f o r m a t i o n of a g i v e n antigen, the an ti b o d i e s gene". in s u pp or t o f views charac te rs a n t i b o d y pro duc tio n. of i n d u c t i o n of m u t a t i o n s have b e e n s u g g e s t e d b y S t u r t e v a n t changes. He has induced by a d v i s e d that there this (194a). res­ is a antigen may cases p r e s e n t e d d e m o n s t r a t i n g the i n h e r i t a n c e of a c q u i r e d s h o u l d be r e e x a m i n e d w i t h the g e n e - a n t i g e n - a n t i b o d y i n t e r a c t i o n p o s s i b i l i t y in mind. The a b o v e m e n t i o n e d considerations make the task of a n ­ alysis of the v e g e t a t i v e h y b r i d i z a t i o n c o n c e p t v e r y i n t r i ­ cate. However, repeat the e x p e r i m e n t s there is no ot he r convincing way than to with all p o s s i b l e m o d i f i c a t i o n s , u nd er -59- st ri c t l y c o n t r o l l e d conditi ons , de fi ni te c o nc lu si on s. in order to arrive at S U M M A R Y -tt-MD CONCLUSIONS The re s u l t s tomatoes claims are of' s t ud ie s r e p o r t e d m a i n l y w i t h the of R u s s i a n s c i e n t i s t s g en et ic are they h a v e d e m o n s t r a t e d the p r e s e n t e d of studies, scion r e l a t i o n s h i p s , relation to to tal f o r m a ti ve 1. ted for that st im ul u s S e v e n graft the study, observed. the No and titrat abl e combinations of the d u r i n g 1947. of the o t h e r g r a f t in v i s i b l e characters which might graft s y m b i o n t s u n d e r solids, titratable any changes produced, and scion, .analyses in f ru it co m p o s i t i o n . were o b s e r v e d w i t h r e g a r d to were fruits i n di ca te of from d i d not On l y v a r i e t a l of an i n t e r ­ for t ot al acid, s ym ­ of lea ve s of fruits -different t r e a t m e n t s , acidity and ascorbic graft of the selec­ on the symbiont, an ot he r va ri e t y , st oc k and six v a r i e t i e s of one the i n f l u e n c e ac t i o n b e t w e e n its ac idity, The e f f e c t s a variety which developed under were stock- ascorb ic acid, of d e f o l i a t e d shoo ts influence ch a n g e s of to i n d u c e d b y 2,4-D. we re m a d e d e v e l o p m e n t of fru it s biont under solids In addition, w i t h r e f er en ce on the s y n t h e s i s s ol ub le in a i m of a n a l y z i n g rece nt i n t e r a c t i o n betvveen s t o c k and scion. the results on the on s t o c k - s e i o n re l a t i o n s h i p s solubl e reveal differences the c o m p o s i t i o n f a cto rs u n d e r study. attempts were made d u r i n g 1948 w i t h the i n t e r a c t i o n of v e r y y o u n g scions the o b j e c t of studying and c o m p a r a t i v e l y old er -61- stocks. Scions w i t h two were s u c c e s s f u l l y leaves cotyledons grafted on and two small leaves s i x - w e e k old consistently developed stocks. abnormalities The scion and studies co ul d not be continued, 2. made Studies on r e c i p r o c a l in their c a p a c it y on the a s c o r b i c gr a f t s to m o n s t r a t e d that the of developed under variety, the was not was content Wo fruits influence of the le av e s c h a n g e d ap pr e c i a b l y . Wo asc or bi c 5. total in the le ave s, so lu bl e a c i d conten t. acidity The ox y a c e t i c de fi n i te the fruits, conclusion synthesized the fruits, the or was s u b st ra te b e i n g the l eav es . stock w i t h o u t t itr ata ble system. of the ot her aci d was tot al soluble solids of fruits which, d e v e l o p e d e i t h e r u n d e r asc orbic root of one v a r i e t y w h i c h and t h e n t r a n s l o c a t e d into changes The de­ acid p r o c e e d e d in i n d e p e n d e n t l y of the of i n d e p e n d e n t l y s y n t h e s i z e d in s u pp li ed by It was obse rv ed. could be d r a w n -as to w h e t h e r in the lea ve s were differing greatly acid. f o r m a t i o n of a s c o r b i c Wo r o o t s t o c k i n f l u e n c e aci d conte nt of fruits two v a r i e t i e s synthesize ascorbic the g r e e n a s s i m i l a t i n g p ar ts ascorbic acid acid was in f l u e n c e or r o o t s t o c k w i t h leaves, so li d s Wo r e l a t i o n s h i p was stimulus transmitted of ro ot - were no ti ced . s h ow e d a r e l a t i o n s h i p to and a s c o r b i c formative the and ac i d co ntents the fo un d b e t w e e n the acid. i n d u c e d by 2 , 4 - d i c h l o r o p h e n through graft unions of tomato - 62- plants from stock to ch em ic al n a t u r e but 4„ The resul ts for a d d i t i o n a l hybridization p o r t e d ha v e clusions „ work scion. its The actual of the stimulus form was not determined,, investigations, to a n a l y z e more concept, h a v e s e e m e d to be of w i t h s u gg est ion s thoroughly, b e e n d is cu ss ed . been considered inadequate for the v e g e t a t i v e The results- r e ­ d r a w i n g final c on ­ BIBLIOGRAPHY Ashby, A. 1946. 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