E F F E C T O F S T O C K - S C I O N I N T E R R E L A T I O N S H I P S AN D G R A F T UNIONS
U PON N U T R I E N T ABSOR PTION A ND T R A N S P O R T IN H I G H E R P L A N T S
AS I N D I C A T E D BY R A D I O A C T I V E IS O T O PE S

By
M A R T I N JOHN BUKOVAC

AN ABSTRACT
S u b m i t t e d to t h e Sc ho ol f o r A d v a n c e d G r a d u a t e S t u d i e s of M i c h i g a n
S t a t e U n i v e r s i t y of A g r i c u l t u r e a n d A p p l i e d S c i e n c e
in p a r t i a l f u l f i l l m e n t of t h e r e q u i r e m e n t s
f o r t h e d e g r e e of
D O C T O R O F PH IL O SO PH Y
D e p a r t m e n t of H o r t i c u l t u r e

M A R T I N JOHN BUKOVAC

ABSTRACT

B ud ded a p p l e a n d v a r i o u s l y g r a f t e d t o m a t o p l a n t s , g r o w n in s o l u t i o n
a n d s o i l c u l t u r e s , w e r e u s e d to d e t e r m i n e t h e e f f e c t of th e g r a f t u n i o n a n d
s t o c k - s c i o n i n t e r r e l a t i o n s h i p s upon f o l i a r a n d r o o t a b s o r p t i o n of n u t r i e n t s
and th e ir subsequent distribution.

R adioactive a n a ly s is and a u to ra d io g ra p h y

of p l a n t t i s s u e s w e r e e m p l o y e d to a s c e r t a i n a b s o r p t i o n a n d s u b s e q u e n t d i s ­
t r i b u t i o n of i s o t o p i c a l l y l a b e l e d n u t r i e n t s .

R o o t e d c u t t i n g s of E a s t M a i l i n g

(EM) VII a n d IX ( d w a r f a p p l e r o o t s t o c k s ) a b s o r b e d l e s s P
r o o t m e d i u m , t h a n E M XVI, a v i g o r o u s a p p l e r o o t s t o c k .

32

, a p p l i e d to t h e

I r r e s p e c t i v e of

f o l i a r o r r o o t a p p l i c a t i o n s , M c I n t o s h a p p l e s c i o n s on E M VII o r IX r o o t s to c k s (dwarf) a b s o r b e d and a c c u m u la te d l e s s P

32

XVI o r D e l i c i o u s s e e d l i n g ( v i g o r o u s ) r o o t s t o c k s .

t h a n w h e n b u d d e d on E M
S i m i l a r l y , l e s s Ca

45

was

r e c o v e r e d in t h e M c I n t o s h s c i o n 144 h o u r s f o l l o w i n g t r e a t m e n t , w h e n a p p l i e d
to t h e m e d i u m in w h i c h r o o t s of M c I n t o s h s c i o n s on E M VII o r IX w e r e
g r o w i n g t h a n in s c i o n s on E M XVI o r D e l i c i o u s s e e d l i n g r o o t s t o c k s .
m a r k e d a c c u m u l a t i o n of P

32

o r Ca

45

No

w a s n o t e d a t t h e bud unio n.

G e n e t i c a l l y d w a r f ( v a r . M a c A r t h u r 741) a n d n o n - d w a r f ( v a r . S t o k e s d a l e ) t o m a t o p l a n t s a b s o r b e d e q u a l q u a n t i t i e s of f o l i a r a p p l i e d P
l e s s of t h e a b s o r b e d P

0 9

32

w a s t r a n s p o r t e d to t h e r o o t s o f t h e d w a r f .

, however,
Tom ato

p l a n t s (v a r. S tokesdale) c h e m i c a l l y d w a rfe d with m a l e i c h y d r a z id e a b s o r b e d
le s s fo liar applied P

32

t h a n n o n - d w a r f e d , but no d i f f e r e n c e s w e r e o b s e r v e d

in t h e s u b s e q u e n t d i s t r i b u t i o n of t h e a b s o r b e d i s o t o p e .
significant d iffe re n c e s o c c u r r e d between the P

32

A s in t h e a p p l e , no

r e c o v e r e d in th e g r a f t u n io n

o r a c o m p a r a b l e s t e m s e g m e n t on a n o n - g r a f t e d t o m a t o p l a n t f o l l o w i n g a b s o r p t i o n of P

32

by t h e r o o t s .

L i k e w i s e , no d i f f e r e n c e s w e r e o b s e r v e d in t h e P

a c c u m u l a t e d in t h e s c i o n s of g r a f t e d a n d n o n - g r a f t e d t o m a t o p l a n t s
48 a n d 96 h o u r s a f t e r a p p l y i n g P

32

to th e r o o t m e d i u m .

s e , d i d not i m p e d e t h e t r a n s p o r t of r o o t a b s o r b e d P

32

.

3

,

1 2

32

, 24,

T h e g r a f t union, p e r
Well f o r m e d g ra ft

u n i o n s m t o m a t o s t e m s did not i m p e d e th e a c r o p e t a l t r a n s p o r t o f Na

99

, S

35

,

^36
_ 45
52-54 ^ 55-59
64
65
86
o 89
99
140
Cl , C a , M n
, Fe
, Cu , Z n , R b , S r , Mo
a n d Ba
f o l l o w i n g r o o t a b s o r p t i o n of t h e i s o t o p e s .

H o w e v e r , w h e n l a t e r a l s of v i g o r -

o u s t o m a t o p l a n t s ( v a r . S t o k e s d a l e ) w e r e g r a f t e d " a p e x to a p e x " P 32 t r a n s p o r t
w a s s i g n i f i c a n t l y r e d u c e d a n d C a 45 t r a n s p o r t w a s a l m o s t c o m p l e t e l y b lo c k e d ,
f o l l o w i n g t h e a b s o r p t i o n of t h e i s o t o p e by t h e r o o t s of o ne of t h e g r a f t e d p l a n t s .
W h e n two o n e - c e n t i m e t e r p h l o e m s e g m e n t s w e r e r e m o v e d f r o m t h e
s t e m s of y e a r o l d a p p l e t r e e s (E M I) a n d r e p l a c e d in a n i n v e r t e d o r n a t u r a l
positio n , m o r e fo lia r a p p lie d P^ 2 a c c u m u la te d above the in v e r s io n s than
above n o n -in v e rte d rings.

L e s s e r a m o u n t s of f o l i a r a p p l i e d P

above the n o n - in v e r te d rin g s.

32

accum ulated

A p p r o x i m a t e l y 60 p e r c e n t m o r e a d v e n t i t i o u s

buds g re w below the in v e r te d than n o n -in v e rte d phloem rin g s.

Swellings

i m m e d i a t e l y a b o v e t h e u p p e r p h l o e m r i n g a n d r e d u c e d top g r o w t h w e r e e v i ­
d e n t o n t r e e s w i t h i n v e r t e d c o m p a r e d to t r e e s w it h n o n - i n v e r t e d p h l o e m

rings.

G r o w t h of th e s c i o n in t e r m s of d r y w e i g h t a n d t e r m i n a l g r o w t h w a s

s i g n i f i c a n t l y i n f l u e n c e d by i t s o r i e n t a t i o n .

L e s s r a d i o c a l c i u m a b s o r b e d by

t h e r o o t s a c c u m u l a t e d in th e i n v e r t e d s c i o n tw o o r s i x d a y s f o l l o w i n g t r e a t ­
m e n t th an with n o n - i n v e r t e d o r n o n - g r a f t e d p la n ts .

H o w e v e r , w h e n th e

c a l c i u m c o n t e n t w a s e x p r e s s e d a s c o u n t s p e r m i n u t e p e r g r a m d r y w ei g h t,
no s i g n i f i c a n t d i f f e r e n c e s w e r e o b s e r v e d , t h u s i n d i c a t i n g t h a t g r o w t h a n d
not th e u n i o n w a s th e d e t e r m i n i n g f a c t o r .

In c o n t r a s t to t h e m a n y e x p e r i -

m e n t s i l l u s t r a t i n g th e a b s e n c e of d o w n w a r d t r a n s p o r t of f o l i a r a p p l i e d C a
a n e s t h e t i z e d b e a n a n d r a d i s h l e a v e s fo l l o w i n g t r e a t m e n t w i t h Ca

45

45

trans­

p o r t e d c o n s i d e r a b l e q u a n t i t i e s out of th e t r e a t e d l e a f to a l l o t h e r p a r t s of
the plan t.

C h e l a t i o n of r a d i o c a l c i u m w i t h d i s o d i u m e t h y l e n e d i a m i n e t e t r a ­

a c e t a t e , p i c r o l i n i c a c id , 2, 5 - d i c h l o r o - 3 ,

6

-dihydroxy-p-quinone and 1-am ino-

2 - n a p t h o l - 4 - s u l f o n i c a c i d d id not e f f e c t i t s b a s i p e t a l t r a n s p o r t .

E F F E C T O F S T O C K - S C I O N I N T E R R E L A T I O N S H I P S A N D G R A F T UNIONS
UPON N U T R I E N T ABSO RPT ION AN D T R A N S P O R T IN H IG H E R P L A N T S
AS I N D I C A T E D BY R A D I O A C T I V E IS OTO PES

By

M A R T I N JOHN BUKOVAC

A THESIS
S u b m i t t e d to t h e S c h o o l f o r A d v a n c e d G r a d u a t e S t u d i e s of M i c h i g a n
S t a t e U n i v e r s i t y of A g r i c u l t u r e a n d A p p l i e d S c i e n c e
in p a r t i a l f u l f i l l m e n t of t h e r e q u i r e m e n t s
f o r t h e d e g r e e of
D O C T O R O F PH IL O SO PH Y
D e p a r t m e n t of H o r t i c u l t u r e

1957

ProQuest Number: 10008270

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TO

MY

PARENTS

John a n d S a d i e Bukovac

ACKNOW LEDGM ENTS

T h e a u t h o r w i s h e s to e x p r e s s h i s s i n c e r e a p p r e c i a t i o n to
D r s . S. H. W i t t w e r a n d H. B. T u k e y f o r t h e i r g u i d a n c e a n d e n c o u r ­
a g e m e n t d u r i n g t h e c o u r s e of th e i n v e s t i g a t i o n a n d p r e p a r a t i o n of
this m a n u s c rip t.
A p p r e c i a t i o n is a l s o e x t e n d e d to m e m b e r s of m y c o m m i t t e e :
D r s . A. L. K e n w o r t h y , R. S. B a n d u r s k i , a n d R. M. S w e n s o n f o r
t h e i r h e l p f u l s u g g e s t i o n s a n d e d i t i n g of t h i s m a n u s c r i p t .
T h e i n t e r e s t a n d t e c h n i c a l a s s i s t a n c e of D r s .

F. G. T e u b n e r

a n d W. G. L o n g a r e g r a t e f u l l y a c k n o w l e d g e d .
T h e w r i t e r d e e p l y a p p r e c i a t e s t h e c o o p e r a t i o n of th e B i o lo g ic a l
a n d M e d i c a l D i v i s i o n of t h e A t o m i c E n e r g y C o m m i s s i o n u n d e r w h o s e
financial a s s i s t a n c e this w o rk was m a d e p o ssib le .

TA BLE OF CO N TE N TS
Page
I N T R O D U C T I O N ..............................................................................................................

1

R E V I E W O F L I T E R A T U R E .....................................................................................
S t o c k - S c i o n I n t e r r e l a t i o n s h i p s ......................................................................
G e n e r a l c o n s i d e r a t i o n s ...........................................................................
S t o c k - s c i o n i n t e r a c t i o n s ......................................................................
A n a t o m i c a l a s p e c t s of s t o c k - s c i o n r e l a t i o n s h i p s ....................
T h e r o l e of th e g r a f t u n io n in s t o c k - s c i o n i n t e r r e l a t i o n s h i p s - e f f e c t o n n u t r i e n t t r a n s p o r t .................................................................
B i o c h e m i c a l s p e c i f i c i t y ...........................................................................
R o o t s t o c k e f f e c t on n u t r i e n t a b s o r p t i o n a n d a c c u m u l a t i o n .
T r a n s l o c a t i o n S t u d i e s .........................................................................................
G e n e r a l c o n s i d e r a t i o n s on t r a n s p o r t .............................................
T r a n s p o r t of p h o s p h o r u s ...................... „ ..........................................
T r a n s p o r t of c a l c i u m ...............................................................................
T r a n s p o r t a n d p o l a r i t y ..........................................................................

2
2
2
3

7
10
13
18
18
20
20
22

G E N E R A L M E T H O D S AN D M A T E R I A L S .......................................................
P r e p a r a t i o n of A p p l e S t i o n s
........................................................
G r o w i n g P l a n t s ......................................................................................................
I s o t o p e S t u d i e s ......................................................................................................

23
23
23
24

E X P E R I M E N T A L R E S U L T S ....................................................................................
S e c t i o n A: E f f e c t of S t o c k - S c i o n I n t e r r e l a t i o n s h i p upon N u t r i e n t
A b s o r p t i o n a n d T r a n s p o r t ..........................................................................
E x p e r i m e n t 1 ..........................
E x p e r i m e n t 2 ................................................................................................
E x p e r i m e n t 3 ................................................................................................
E x p e r i m e n t 4 ................................................................................................
E x p e r i m e n t 5 ................................................................................................
E x p e r i m e n t 6 ................................................................................................
S e c t i o n B: E f f e c t of t h e G r a f t U nion a n d S c io n O r i e n t a t i o n on
N u t r i e n t T r a n s p o r t ......................................................................................
E x p e r i m e n t 7 ...............................................................................................
E x p e r i m e n t 8 ...............................................................................................
E x p e r i m e n t 9 ...............................................................................................
E x p e r i m e n t 1 0 ...........................................................................................
E x p e r i m e n t 1 1 ...........................................................................................
E x p e r i m e n t 1 2 ...........................................................................................

29

6

29
29
31
31
31
45
47
50
50
53
54
59
67
79

S e c t i o n C: S o m e F a c t o r s A f f e c t i n g T r a n s p o r t in t h e P h l o e m . . .
E x p e r i m e n t 1 3 .....................................................................................................
E x p e r i m e n t 1 4 .....................................................................................................

86

D I S C U S S I O N ..............................................................................................................................

91

S U M M A R Y .................................................................................................................................

100

G L O S S A R Y .................................................................................................................................

103

L I T E R A T U R E C I T E D ........................................................................................................

104

86

90

LIST OF TABLES
TABLE

PAGE

I. L E N G T H O F T E R M I N A L G R O W T H O F M C I N T O S H SCIONS F O U R AND
SIX W E E K S A F T E R BUD BREAK ON D W A R F I N G A N D VIGOROUS
R O O T S T O C K S R E C O R D E D IN T H R E E E X P E R I M E N T S ..............................

36

II. R A D I O A C T I V I T Y IN BUD UNIONS 96 HO URS F O L L O W I N G F O L I A R A P ­
P L I C A T I O N O F P 3 2 A N D R O O T A P P L IC A T IO N S OF P 3 2 AN D C a 4 5 . .

39

III.

IV.

V.

VI.

VII.

VIII.

IX.

T R A N S P O R T O F RADIO PHOSPH ORU S <P32) A N D R A D I O C A L C I U M ( C a 4 5 )
IN M C I N T O S H A P P L E SCIONS BUDDED ON E A S T M A L L I N G VII, IX
A N D XVI A N D D E L I C I O U S S E E D L I N G R O O T S T O C K S ...................................

40

ABSO RPT ION AND T R A N S L O C A T I O N O F F O L I A R A P P L I E D R A D I O ­
PHOSPHORUS (P32) BY G E N E T I C A L L Y D W A R F ( M A C A R T H U R 741)
AN D N O N - D W A R F (S T O K E S D A L E ) T O M A T O P L A N T S ..............................

46

T R A N S P O R T O F F O L I A R A P P L IE D RADIOPHOSPHORUS (P32) IN
C H E M I C A L L Y D W A R F E D T O M A T O P L A N T S (VAR. S T O K E S D A L E ) .

49

E F F E C T O F T H E G R A F T UNION ON A C R O P E T A L T R A N S P O R T OF
p32 i n T O M A T O (VAR. S T O K E S D A L E ) S T E M S ..............................................

52

T R A N S P O R T O F P 3 2 A N D C a 4 5 A C RO S S G R A F T UNIONS (APEX T O
APEX) INVOLVING R E V E R S E D P O L A R I T Y IN T H E T O M A T O . . . .

64

G R O W T H O F SCION ( T O M A T O / T O M A T O ) AS I N F L U E N C E D BY
O R I E N T A T I O N O F S C I O N ..............................................................................................

72

G R O W T H O F SCION ( T O M A T O / T O M A T O ) AS I N F L U E N C E D BY
O R I E N T A T I O N O F B U D ...................................................................................................

74

X. T O T A L R O O T ABSORBED R A D I O C A L C I U M ( C a 4 5 ) R E C O V E R E D IN
T H E SCI O N AN D G R A F T UNIONS O F V A R I O U S L Y G R A F T E D T O M A T O
P L A N T S ......................................................................................................................................

75

45
R O O T ABSORBED R A D I O C A L C I U M (Ca ) R E C O V E R E D IN T H E SCION
A N D G R A F T UNIONS O F V A R I O U S L Y G R A F T E D T O M A T O PLA NTS .

76

XI.

TABLE
XII.

XIII.

XIV.

XV.

Page
T O T A L R O O T ABSORBED R A D I O C A L C I U M ( C a 4 5 ) R E C O V E R E D IN
T H E SCIONS AND BUD UNIONS O F V A R I O U S L Y BUDDED T O M A T O
P L A N T S .................................................................................................................................

77

R O O T ABSORBED R A D I O C A L C I U M ( C a 4 5 ) R E C O V E R E D IN T H E
SCIO N AND BUD UNIONS O F V A R I O U S L Y BUDDED T O M A T O P L A N T S

78

N U MB ER O F BUDS F O R C E D B E L O W T W O P H L O E M RINGS ON A P P L E
A N D T O B A C C O P L A N T S AS A R E S U L T OF R E M O V I N G , IN V E R T IN G
AND R E P L A C I N G , R E M O V I N G AND N O T R E P L A C I N G , OR R E M O V ­
ING AN D R E P L A C I N G IN T H E N O R M A L P O S IT IO N T W O RINGS OF
P H L O E M .................................................................................................................................

81

S I Z E O F S T E M S W E L L I N G AND T E R M I N A L G R O W T H O F O N E - Y E A R O L D A P P L E T R E E S ( E A S T M A L L I N G I) AS I N F L U E N C E D BY R E M O V ­
AL AND IN V E R SI O N OR R E M O V A L AND N O R M A L R E P L A C E M E N T O F
T W O P H L O E M R I N G S .....................................................................................................
82

L IS T O F FIG U RES
F igure

Page
32
A c c u m u l a t i o n of r a d i o p h o s p h o r u s (P ) in t h e t o p s of r o o t e d c u t ­
t i n g s of E a s t M a i l i n g (EM) VII, IX a n d XVI, 24, 48 a n d 96 h o u r s
a f t e r a p p l y i n g P^ 2 to t h e r o o t m e d i u m .......................................................

30

2.

S o l u t i o n c u l t u r e s u s e d in g r o w i n g t h e p l a n t s ..........................................

32

3.

R a d i o p h o s p h o r u s (p32 c p m / g m d r y w t . ) r e c o v e r e d in t h e v a r i o u s
t i s s u e s a t 24, 48 a n d 96 h o u r s f o l l o w i n g f o l i a r a p p l i c a t i o n of P ^
l a b e l e d o r t h o - p h o s p h o r i c a c i d (0. 2 %) to M c I n t o s h s c i o n s b u d d e d
o n E a s t M a i l i n g (EM) VII, IX a n d XVI a n d D e l i c i o u s s e e d l i n g r o o t ­
s t o c k s ............................................................................................................................

37

R a d i o p h o s p h o r u s (P 32 c p m / g m d r y w t . ) r e c o v e r e d in v a r i o u s
t i s s u e s of M c I n t o s h s c i o n s b u d d e d on E a s t M a i l i n g (EM) VII, IX
a n d XVI a n d D e l i c i o u s s e e d l i n g r o o t s t o c k s , 24, 48 a n d 96 h o u r s
22
f o l l o w i n g t r e a t m e n t w ith P
...........................................................................

41

45
R a d i o c a l c i u m (Ca
c p m / g m d r y w t . ) d e t e c t e d in t h e M c I n t o s h
s c i o n s on E M VII, IX a n d XVI r o o t s t o c k s , 48, 96 a n d 144 h o u r s
f o l l o w i n g i t s a d d i t i o n to t h e r o o t m e d i u m ..............................................

44

A u t o r a d i o g r a m s of g r a f t e d t o m a t o s t e m s s h o w i n g d i s t r i b u t i o n of
C a 4 5 , Ba-*-4 ^, a n d S r ^ 9 t 48 h o u r s f o l l o w i n g t h e a d d i t i o n of th e
i s o t o p e s to t h e r o o t m e d i u m ..........................................................................

55

A u t o r a d i o g r a m s o f g r a f t e d t o m a t o s t e m s s h o w i n g d i s t r i b u t i o n of
C i 3 6 } g35 ancj j\ja 2 2 j 4 3 h o u r s f o l l o w i n g t h e a d d i t i o n of t h e i s o t o p e s
to t h e r o o t m e d i u m ..............................................................................................

56

A u t o r a d i o g r a m s of g r a f t e d t o m a t o s t e m s i l l u s t r a t i n g t h e d i s t r i ­
b u t i o n o f P ^ , R b ^ 6 anci M n 5 2 - 5 4 j 4 8 h o u r s f o l l o w i n g s o i l a p p l i ­
c a t i o n ...........................................................................................................................

57

A u t o r a d i o g r a m s of g r a f t e d t o m a t o s t e m s i l l u s t r a t i n g t h e d i s t r i ­
b u t i o n of Z n ^ a n d M o ^ , 48 h o u r s a f t e r t r e a t i n g t h e r o o t m e d i u m

58

1.

4.

5.

6

.

7.

8

.

9.

F ig u re
10.

11.

12.

13.

14.

15.

Page
P h o t o g r a p h i l l u s t r a t i n g a l o n g i t u d i n a l s e c t i o n t h r o u g h a C l a r k ’s
d w a rf i n t e r s t e m w ith a M cIn to sh scio n and H ib e rn a l stock, and
a u t o r a d i o g r a m s h o w i n g th e d i s t r i b u t i o n of C a ^ , 24 h o u r s a f t e r
a p p l i c a t i o n to t h e r o o t m e d i u m ...........................................................................

60

A u n i o n f o r m e d by g r a f t i n g t o g e t h e r ( a p e x to a p e x ) t h e f o r c e d
l a t e r a l s of tw o t o m a t o p l a n t s ................................................................................

61

P h o t o g r a p h a n d a u t o r a d i o g r a m of a n " a p e x to a p e x " g r a f t u n io n
b e t w e e n a l a t e r a l f r o m o ne t o m a t o p l a n t a n d t h a t of a n o t h e r . . .

63

P h o t o g r a p h s a n d a u t o r a d i o g r a m s of i n a r c h e d s t e m s of t o m a t o
p l a n t s s h o w i n g t h e d i s t r i b u t i o n of r o o t a b s o r b e d r a d i o c a l c i u m
...................................................................
(C a ^ 5 ) 48 h o u r s a f t e r t r e a t m e n t

66

P h o t o g r a p h s s h o w i n g g r o w t h of s c i o n s on d e c a p i t a t e d t o m a t o p l a n t s
2 1 d a y s a f t e r b u d d i n g a n d 1 1 d a y s a f t e r g r a f t i n g ......................................

68

A u t o r a d i o g r a m s i l l u s t r a t i n g t h e d i s t r i b u t i o n of r a d i o c a l c i u m a s i n ­
f l u e n c e d by s c i o n o r i e n t a t i o n in b u d d e d a n d g r a f t e d t o m a t o p l a n t s ,
48 h o u r s f o l l o w i n g a p p l i c a t i o n of C a ^ to t h e r o o t m e d i u m . . . .

71

16.

P h o to g r a p h s show ing the c o m p a r a t i v e q u a n titie s ( c p m / g m d ry w t . )
of f o l i a r a p p l i e d r a d i o p h o s p h o r u s r e c o v e r e d in th e v a r i o u s p h l o e m
s e g m e n t s in t h e a p p l e a n d r e g e n e r a t i o n of th e x y l e m in th e v i c i n i t y
of t h e p h l o e m r i n g s ..................................................................................
84

17.

P h o t o g r a p h s a n d a u t o r a d i o g r a m s s h o w i n g t h e d i s t r i b u t i o n of f o l i a r
applied P ^ through in v e rte d and n o n -in v e rte d phloem rin g s . . .

18.

19.

A u t o r a d i o g r a m s s h o w i n g t h e d i s t r i b u t i o n of f o l i a r a p p l i e d C a
th e b e a n p l a n t ..............................................................................................

45

in
88

P h o t o g r a p h s a n d a u t o r a d i o g r a m s of m o u n t e d r a d i s h h y p o c o t y l s p e c i ­
m e n s s h o w i n g d i s t r i b u t i o n of f o l i a r a p p l i e d C a ^ f o l l o w i n g a n e s t h e t i ­
z a t i o n of t h e C a ^ t r e a t e d l e a f .........................................................
89

85

I. IN TR O D U C TIO N

T h e a r t of g r a f tin g h a s b e e n know n to m a n k in d s in c e th e e r a b e fo re
C h r is t, y e t th e i n t r i c a t e r e la tio n s h ip s b e tw e e n s to c k an d s c io n r e m a in o b s c u r e .
T h e d w a rf r e s p o n s e o f s c io n to r o o ts to c k is o n ly one of th e m a n y c o m p le x
m a n if e s ta tio n s of s to c k - s c io n in te r r e la tio n s h ip .

R o o ts to c k e ffe c t upon sc io n ,

s c io n on r o o ts to c k a n d i n te r s te m on both, w ith th e i r m a n y m a r k e d p h y s io lo g ic a l
a n d m o rp h o lo g ic a l c h a n g e s , h av e in tr ig u e d th e s c ie n tif ic m in d f o r m a n y y e a r s .
In th e p a s t d e c a d e o r two, th e A m e r ic a n h o r ti c u l t u r i s t h a s b e c o m e e x ­
t r e m e l y i n t e r e s t e d in u tiliz in g s o m e of th e s e p h y s io lo g ic a l r e s p o n s e s to c o n ­
t r o l th e p ro d u c tio n of c o m m e r c ia l f r u it c r o p s .

As an e x a m p le , th e n e e d s of a

r a p id ly e x p a n d in g h o r ti c u l t u r a l in d u s tr y fo r u n ifo rm , e a r l i e r fru itin g , d w a rf
o r s e m i- d w a r f a p p le t r e e s h a s m a te r ia lly a d d e d to th e p o p u la r ity of the
1
M a ilin g r o o ts to c k s .
In v iew of th is i n c r e a s e d d e m a n d th e n e e d f o r a m o re c o m p le te u n d e r ­
s ta n d in g of s to c k - s c io n r e la tio n s h ip h a s a c q u ir e d a g r e a t e r im p o rta n c e .

The

w o rk w h ich fo llo w s w as u n d e rta k e n , w ith th e a id of r a d io a c tiv e is o to p e s , to
a s c e r t a i n th e n u tr itio n a l r o le of th e g r a f t u n io n an d r o o ts to c k in th e p h y s io lo g y
o f d w a rfin g .

S p e c ia l c o n s id e r a tio n h a s b e e n g iv en to a b s o r p tio n a n d s u b s e ­

q u en t t r a n s p o r t o f e s s e n t i a l n u tr ie n ts .
*A p p le c lo n e s o r ig in a tin g fro m th e P a r a d is e o r D oucin g ro u p s of a p p le s
( c l a s s i f i e d a s M a lu s s y lv e s t r i s , M ill, by B ailey 1949) w e r e u s e d a s r o o ts to c k s
in a g r a f t c o m b in a tio n sh o w in g v a r io u s d e g r e e s of d w a rfis m on th e g ro w th of
th e s c io n ; h e r e a f t e r , sy n o n y m o u s w ith E a s t M a ilin g o r E . M.

II. R E V I E W O F L I T E R A T U R E
Stock-Scion In te rre la tio n s h ip s

G e n era l considerations:

It i s n o t u n u s u a l t h a t c o m p l e x i n t e r a c t i o n s

a r i s e a s a r e s u l t of g r a f t i n g , w h e n o n e c o n s i d e r s t h a t tw o s e p a r a t e i n d e p e n d e n t
in d iv id u als a r e b rought to g e th e r, w hich unite a n d g ro w a s one.

These in te r­

a c t i o n s a r e of g r e a t p r a c t i c a l i m p o r t a n c e to th e f r u i t g r o w e r a n d h a v e b e e n
re a liz e d as such for m any y ears.

A s e a r l y a s 1818 T h o m a s A n d r e w K ni ght

(1818) r e p o r t e d t h a t p l u m s t o c k s w e r e b e s t a d a p t e d to t h e p e a c h , a n d a p r i c o t
s t o c k s to M o o r ’s P a r k a p r i c o t .

F u r t h e r m o r e , he n o t e d a s m o o t h h ig h l y

p o l i s h e d b a r k a n d a h e a l t h i e r m o r e v i g o r o u s t r e e w h e n M o o r ’s P a r k w a s
b u d d e d on a p r i c o t s t o c k t h a n on a n y o t h e r r o o t s t o c k .
M a n y g r a f t i n g f a i l u r e s h a v e b e e n a t t r i b u t e d to s t o c k - s c i o n i n c o m p a t i ­
bility.

A t t e m p t s h a v e b e e n m a d e to e x p l a i n i n c o m p a t i b i l i t y on t h e b a s i s of

a n a t o m i c a l d i f f e r e n c e s ( D a n i e l 1894, G u i g n a r d 1907, C r a f t s 1934,

Beakbane

a n d T h o m p s o n 1939, a n d B e a k b a n e 1941) a n d d e f e c t i v e g r a f t u n i o n s (W a ugh 1904
a n d B r a d f o r d a n d S i t t o n 1926).

B i o c h e m i c a l d i f f e r e n c e s h a v e b e e n s u g g e s t e d by

G r e e n (1926), M i l b r a i t h a n d Z e l l e r (1945), a n d De S t i g t e r (1956), a n d a n a n t i ­
g e n a n t i b o d y p h e n o m e n o n h a s b e e n p r o p o s e d by K o s t o f f (1928,

1929,

1931).

V y v y a n (1934) s u g g e s t e d a d i f f e r e n t i a l g r o w t h r a t e b e t w e e n s t o c k a n d s c i o n ,
w h e r e a s S w a r b r i c k e t a l (1946) s u g g e s t e d a d i f f e r e n t i a l a b s o r p t i o n a n d

3

s u b s e q u e n t t r a n s p o r t o r a c o m b i n a t i o n of th e tw o a s c o n t r i b u t i n g f a c t o r s in
s to c k -sc io n incom patibility.
Stock-scion interactions:

T h e p h y s i o l o g i c a l e f f e c t of s t o c k on s c i o n o r

s c i o n on s t o c k i s n o t l o c a l i z e d in a n y o n e c o m p o n e n t of a g r a f t e d c o m b i n a t i o n .
H a t t o n (1927) s t u d i e d t h e e f f e c t of a n u m b e r of a p p l e r o o t s t o c k s upon t h e s c i o n
a n d c o n c l u d e d t h a t g r o w t h , v i g o r a n d f r u i t f u l n e s s of t h e s c i o n w a s i n f l u e n c e d
by r o o t s t o c k .

T r e e s g r o w n on M a i l i n g VIII o r IX w e r e s m a l l a n d u s u a l l y p r o ­

d u c e d s w e l l i n g s a t t h e g r a f t unio n, w h i l e t h o s e g r o w n on M a i l i n g II a n d XVI
w ere m o re vigorous.

A m o s e t a l (1936) c o n c l u d e d a f t e r t w e n t y y e a r s of o b s e r ­

v a t i o n s a t E a s t M a i l i n g , t h a t M a i l i n g VIII a n d IX r o o t s t o c k s r e s u l t e d in d w a r f ­
ing a n d e a r l i e r fr u itin g w as a s s o c i a t e d with the s c io n s g r a f t e d th e r e o n .

The

d w a r f i n g r e s p o n s e of s c i o n to s t o c k w a s o n l y o n e of t h e m a n y m a n i f e s t a t i o n s
observed.

M a i l i n g VII, w h i c h i s c o n s i d e r e d a s e m i - d w a r f r o o t s t o c k , i m p a r t s

v i g o r o u s g r o w t h to t h e s c i o n in t h e e a r l y y e a r s .

H ow ever, the g row th r a t e

f a l l s off o n c e t h e t r e e b e g i n s to f r u i t .
T h a t a s t o c k e f f e c t m a y b e e x e r t e d by a s t e m p i e c e w a s c l e a r l y d e m o n ­
s t r a t e d by S w a r b r i c k a n d R o b e r t s (1927) a n d R o b e r t s (1929).
using various in te rste m s,

T hese w orkers,

s h o w e d t h a t t h e d w a r f i n g e f f e c t of a s t o c k c o u l d be

p r o d u c e d by a s t e m p o r t i o n of th e s t o c k .

T h i s w a s l a t e r c o n f i r m e d by H a t t o n

(1930), V y v y a n (1934), T u k e y a n d B r a s e (1933,
S a x (1953) a n d T h o m a s (1954).

1943) a n d m o r e r e c e n t l y by

In t h e y e a r s t h a t fo ll o w e d , t h e r e l a t i v e

4

i m p o r t a n c e of t h e s t o c k a n d s t o c k - s t e m b e c a m e t h e o b j e c t of e x t e n s i v e e x p e r i ­
m entation.

T u k e y a n d B r a s e (1933) c o n c l u d e d a f t e r e x t e n s i v e i n v e s t i g a t i o n s

t h a t no o n e p a r t d o m i n a t e d t h e e n t i r e p l a n t .

They showed that all had a m a r k e d

influ en ce a t one tim e o r a n o th er, how ever, the r o o ts to c k e x e r t e d the g r e a t e s t
influence.

T h i s f i n d i n g h a s b e e n c o n f i r m e d by K n ig h t (1935) a n d V y v y a n (1938,

1955) who a g r e e d , in p a r t , t h a t t h e r o o t s t o c k e x e r t e d t h e g r e a t e r i n f l u e n c e .
B e a k b a n e , H a t t o n a n d A m o s (1937) s h o w e d t h a t s c i o n s g r a f t e d on a l o n g s t o c k s t e m p i e c e w e r e i n f l u e n c e d to a g r e a t e r d e g r e e by th e r o o t s t o c k t h a n s c i o n s
g r a f t e d d i r e c t l y to a r o o t - p i e c e .

M o r e r e c e n t l y , B e a k b a n e a n d R o g e r s (1956)

c o n c l u d e d t h a t t h e r o o t p o r t i o n s of c l o n a l a p p l e r o o t s t o c k s a r e c a p a b l e of
e x e r t i n g an i n f l u e n c e c h a r a c t e r i s t i c of t h e c l o n e in t h e a b s e n c e of a n y r o o t stock stem .

F u r t h e r m o r e , t h e p r e s e n c e of a s t o c k - s t e m on th e r o o t s t o c k

in te n sifie d the ro o ts to c k effect.
T h a t p h y s i c a l c h a r a c t e r i s t i c s of r o o t s of d w a r f a n d v i g o r o u s a p p l e
t r e e s di d n o t d i f f e r m a t e r i a l l y w a s r e p o r t e d by R o g e r s a n d V y v y a n (1927),
R o g e r s (1 9 3 9 a ) a n d H a t t o n (1935).

A l s o , r o o t s of a d w a r f i n g r o o t s t o c k p e n e ­

t r a t e d t h e s o i l to d e p t h s e q u a l to t h o s e of v i g o r o u s r o o t s t o c k s ( H a t t o n 1935).
R o g e r s (193 9b ) s t u d i e d , in d e t a i l , r o o t g r o w t h a s r e l a t e d to r o o t s t o c k ,
seasonal and clim atic facto rs.

s o il ,

Root grow th c lo se ly followed the above

g r o u n d v i g o r , t h a t i s , M a i l i n g IX l e a s t v i g o r o u s , M a i l i n g I i n t e r m e d i a t e , a n d
M a i l i n g XVI m o s t v i g o r o u s .

No a p p r e c i a b l e d i f f e r e n c e s in a p p e a r a n c e w e r e

a p p a r e n t b e t w e e n r o o t s of M a i l i n g IX, I a n d XVI.
looked s i m i l a r .

R o o t h a i r s of a l l s t o c k s

H ow ever, when s e c o n d a ry th ic k e n in g o c c u r r e d , M ailin g

IX r o o t s s h o w e d a m o r e y e l l o w i s h c o l o r t h a n e i t h e r M a i l i n g I o r XVI.
R o g e rs concluded;
" t h e d i f f e r e n t r o o t s t o c k s , so c o n t r a s t i n g in t h e i r e f f e c t on
g r o w t h a n d c r o p p i n g , s h o w e d no d r a m a t i c c o n t r a s t in a b s o r b ­
in g r o o t t y p e a n d c o n f o r m a t i o n . T h e m o s t s t r i k i n g f e a t u r e
w a s t h e d e g r e e of s i m i l a r i t y b e t w e e n t h e i n d i v i d u a l r o o t s a n d
b e t w e e n t h e i r r e a c t i o n s to p h y s i o l o g i c a l f a c t o r s "
T h e e f f e c t of s t o c k up on th e d e v e l o p m e n t of t h e s c i o n h a s b e e n w e l l
established.

On t h e o t h e r hand, t h e r e is e v i d e n c e t h a t t h e s c i o n m a y i n ­

f l u e n c e t h e d e v e l o p m e n t of t h e s t o c k ( S w a r b r i c k a n d R o b e r t s 1927, R o b e r t s
1929, T u k e y a n d B r a s e 1933, A r g l e s 1937, a n d R u b i n 1955).
T u k e y a n d B r a s e (1933) w r i t e ;
"A m erican horticultural lite ra tu re re c o rd s observations as
e a r l y a s 1846 t h a t t h e c i o n m a y a f f e c t th e d e v e l o p m e n t of t h e
r o o t s t o c k , a n d o t h e r i n s t a n c e s h a v e b e e n r e c o r d e d s i n c e . It
h a s b e e n s h o w n t h a t t h e c i o n m a y a f f e c t t h e r o o t in o t h e r w a y s
t h a n in q u a n t i t y , d i r e c t i o n , a n d f o r m of r o o t s , s u c h a s l o n g e ­
vi ty , h a r d i n e s s , c h e m i c a l c o m p o s i t i o n , a n d c o l o r . . ................. "
R e c e n t l y , V y v y a n (1955) r e p o r t e d t h a t r o o t s t o c k h a s a g r e a t e r i n f l u ­
e n c e on t r e e s i z e t h a n s c i o n ,

however,

c u l a r l y w h e n M a i l i n g XII w a s t h e s c i o n .

scion v a rie ty had som e effect p a r t i ­
O c c a s i o n a l l y one f i n d s t h a t th e

sc io n d o m in a te s the stock, although this s eld o m is the lim i t i n g f a c to r .
H o d g s o n a n d C a m e r o n (1943) w o r k i n g w i t h c i t r u s f o u n d t h a t w h e n th e v i g o r
of t h e s c i o n i s l e s s t h a n t h a t of t h e r o o t s t o c k , i t w a s th e s c i o n w h i c h d e t e r ­
m i n e d t h e t o t a l g r o w t h of t h e t r e e .

6

T h e r e i s no c l e a r c u t g r o w t h r e s p o n s e of s c i o n to s t o c k o r of s t o c k
to s c i o n .

One v a r i e t y u s e d a s a s t o c k m a y r e s p o n d q u i t e d i f f e r e n t l y w h e n

u s e d a s the scion.

A l s o , t h e s o u r c e of th e s c i o n m a y i n f l u e n c e i t s b e h a v i o r

w h e n g r a f t e d on th e s a m e r o o t s t o c k .

T u k e y a n d B r a s e (1944) r e p o r t e d a

d i f f e r e n c e in t h e p e r f o r m a n c e of M c I n t o s h b u d w o o d t a k e n f r o m t h e s a m e
h o r t i c u l t u r a l v a r i e t y w h e n b u d d e d on r o o t s t o c k USDA 227.

T h i s led the

a u t h o r s to s u g g e s t t h a t t h e c o m p a t i b i l i t y o r i n c o m p a t i b i l i t y w a s d ue to s o m e
f a c t o r o t h e r t h e n g e n e t i c , e. g. v i r u s o r p h y s i o l o g i c a l d i s t u r b a n c e .
T h e p h y s i o l o g i c a l a s p e c t of s t o c k - s c i o n e f f e c t h a s b e e n c o n s i d e r e d
by V y v y a n (19 55).

He s t r e s s e d t h a t th e s t e m / r o o t r a t i o i s r e l a t i v e l y c o n s t a n t

w i t h i n a n y o n e s c i o n / r o o t s t o c k c o m b i n a t i o n , a n d i s i n d e p e n d e n t of t r e e
size o r age.

T h i s m e r i t s c o n s i d e r a b l e a t t e n t i o n if we r e c o g n i z e t h a t t h e

s c i o n a n d s t o c k m u s t m u t u a l l y a d j u s t to e a c h o t h e r ' s p h y s i o l o g i c a l p r o c e s s e s .
A n ato m ical a s p e c ts of s to c k - s c io n re la tio n sh ip s:

The internal s t r u c t ­

u r e of t h e d w a r f i n g r o o t s t o c k M a i l i n g IX w a s s t u d i e d in a n a t t e m p t to e x p l a i n
i t s d w a r f i n g e f f e c t (B e a k b a n e a n d R e n w i c k 1936).

They concluded that M ailing

IX h a d s m a l l e r v e s s e l s a n d a h i g h e r p e r c e n t of m e d u l l a r y r a y s t h a n th e
ro o te d scion.

M o r e d e t a i l e d s t u d i e s (B e a k b a n e a n d T h o m p s o n 1939) r e v e a l e d

a c o r r e l a t i o n b e tw een the b a r k / w o o d ra tio a n d v ig o r.
the lo w e r the b a r k /w o o d ra tio .

The g r e a t e r the v ig o r

T h u s , t h e r a t i o of b a r k to w o o d f o r M a i l i n g

IX w a s 3. 1, a n d 1. 0 f o r M a i l i n g VII, a m o r e v i g o r o u s r o o t s t o c k .

M icroscopic

7

o b s e r v a t i o n s of t h e r o o t s r e v e a l e d t h a t t h e d w a r f i n g s t o c k ( M a i l i n g IX) h a d
a g r e a t e r p e r c e n t o f p a r e n c h y m a a n d r a y c e l l s a n d a l o w e r p e r c e n t of f i b e r s
a n d v e s s e l s t h a n t h e v i g o r o u s s t o c k ( M a i l i n g II).

F u r t h e r m o r e , the v e s s e l s

w e r e g e n e r a l l y l a r g e r in M a i l i n g II, h o w e v e r , no c o r r e l a t i o n w a s fo un d b e ­
tween size and vigor.

Beakbane a n d T h o m p so n a tta c h e d sp ec ia l sig n ifican ce

to t h e h i g h e r p e r c e n t of r a y t i s s u e a n d l o w e r p e r c e n t of v e s s e l s in d w a r f i n g
rootstock.

T h e y c o n clu d e d that a l a r g e r v o lu m e is a v a il a b le for c a r b o h y d r a t e

a c c u m u l a t i o n a n d s t o r a g e in t h e d w a r f i n g s t o c k s .

Thus,

r e s u l t i n g in e a r l i e r

f r u i t i n g a n d a r e d u c t i o n in m i n e r a l s a l t t r a n s p o r t t h r o u g h t h e s m a l l e r v e s s e l s ,
T h i s m a y b e r e s p o n s i b l e f o r t h e r e d u c e d g r o w t h o b s e r v e d in t h e s c i o n s g r a f t e d
on d w a r f i n g s t o c k s .

In a m o r e r e c e n t p a p e r , B e a k b a n e (1941) fo u n d l a r g e r q u a n ­

t i t i e s of s t a r c h , f a t s , o i l s , o x i d a s e s a n d p e r o x i d a s e s a s s o c i a t e d w i t h i n c r e a s e d
p a r e n c h y m a c e l l s in the r o o t s t o c k s of M a i l i n g IX t h a n in M a i l i n g II o r XII.
T h e r o l e of th e g r a f t u n i o n in s t o c k - s c i o n r e l a t i o n s h i p - - e f f e c t on
nutrient transport:

T h e u n i t i n g of t h e s c i o n a n d s t o c k to f o r m a g r a f t u n i o n

i n t r o d u c e s a n u m b e r of v a r i a b l e s , a n y of w h i c h a l o n e o r in c o m b i n a t i o n m a y
influence the s to c k - s c io n re la tio n s h ip .

C o n s i d e r i n g th e g r a f t unio n f r o m p u r e l y

a m e c h a n i c a l view point, s t r u c t u r a l w e a k n e s s e s , a n a to m ic a l and p h y sio lo g ic al
d i f f e r e n c e s m a y all m a t e r i a l l y affect the s t o c k - s c i o n re la tio n s h ip .

These

p o s s i b i l i t i e s w e r e r e c o g n i z e d by D a n i e l (1894), W a u g h (1904), B r a d f o r d a n d
S i t t o n (1926) a n d P r o e b s t i n g (1926,

1928).

W a ugh o b s e r v e d t h a t in i m p e r f e c t

8

u n i o n s t h e c o n t i n u i t y of n e w g r o w t h w a s i n t e r r u p t e d by d e p o s i t i o n of l o o s e
s c a r ti s s u e fo r m e d d u rin g the healin g p r o c e s s .
o t h e r han d,

B r a d f o r d a n d Si t to n , on t h e

s t a t e d t h a t d w a r f i n g w a s a s s o c i a t e d w it h p h l o e m d i s c o n t i n u i t y .

T h e y felt the effe ct w a s r e c i p r o c a l , th at is, the sto ck w a s d w a rfe d f i r s t fo r
l a c k of e l a b o r a t e d fo o d s , and, in t u r n , it d w a r f e d t h e s c i o n .

Proebsting

v i s u a l i z e d a l a y e r of p e r i d e r m f o r m i n g b e t w e e n t h e s c i o n a n d s t o c k in th e c a s e
of apple and p e a r .

T h e s e f a c t o r s a l o n e m a y a c c o u n t f o r th e m a n y r e p o r t s

on i m p e d e d t r a n s p o r t t h r o u g h th e g r a f t u n io n ( V o c h t i n g 1894, G u i g n a r d 1907,
C h a n g 1937, W a r n e a n d R a b y 1939, H e r r e r o 1951, a n d G l u s h c h e n k o a n d D r o b k o v
1952).
T h e e f f e c t of th e g r a f t u n i o n on m i n e r a l o r w a t e r t r a n s p o r t h a s not
b e e n f u l l y e v a l u a t e d in i n t a c t p l a n t s .

W a r n e a n d R a b y (1939), u s i n g t e n -

c e n t i m e t e r s e g m e n t s c o n t a i n i n g t h e g r a f t union, m e a s u r e d t h e q u a n t i t y of
w a t e r w h i c h p a s s e d t h r o u g h in t e n m i n u t e s u n d e r a p r e s s u r e of 16 c e n t i m e t e r s
of m e r c u r y .

T h e y fo u n d t h a t u n i o n s in w h i c h M a i l i n g IX w a s o n e of th e c o m ­

p o n e n t s w e r e l e s s e f f i c i e n t in w a t e r c o n d u c t i o n t h a n t h o s e in w h i c h IX w a s
c o m b i n e d w i t h IX, o r in w h i c h II w a s c o m b i n e d w i t h i t s e l f o r a n o t h e r s c i o n .
M o r e d e ta ile d in v e s t ig a ti o n s r e v e a l e d that at the g ra ft union t h e r e w as l e s s
c o n d u c t i n g t i s s u e t h a n in t h e s c i o n o r s t o c k .

T h i s l e d t h e a u t h o r s to s u g ­

g e s t t h a t t h e g r a f t u n io n i n t r o d u c e d an a d d i t i o n a l r e s i s t a n c e to th e flow of
w ater.

C h a n g (1937) r e p o r t e d t h a t t h e r e d u c e d t r a n s p o r t of w a t e r a c r o s s

9

t h e u n i o n a n d t h e a c c u m u l a t i o n of s t a r c h a b o v e th e u n i o n r e s u l t e d f r o m m e c h ­
anical obstruction.

He fo u n d t h a t o n l y o n e - f i f t h a s m u c h w a t e r p a s s e d t h r o u g h

g r a f t u n i o n s of i n c o m p a t i b l e c o m b i n a t i o n s of p e a r a n d q u i n c e a s t h r o u g h c o m ­
patible com binations.

L a u r e n t (1908) c o n c l u d e d t h a t t h e b o u r r e l e t ( c a l l u s o r

p e r i d e r m l a y e r a r i s i n g b e t w e e n t h e s c i o n a n d s t o c k ) m a y be d i f f e r e n t i a l l y
p e r m e a b l e , t h u s p e r m i t t i n g t h e p a s s a g e of c e r t a i n g l u c o s i d e s a n d n o t of
others.

P h o s p h o r u s , on t h e o t h e r hand, w a s fo und ( G l u s h c h e n k o a n d D r o b k o v

1952) u n e v e n l y d i s t r i b u t e d w i t h i n g r a f t c o m b i n a t i o n s of t o m a t o a n d n i g h t s h a d e .
S o m e a c c u m u l a t i o n of p h o s p h o r u s o c c u r r e d a t the g r a f t un io n .
In c o n t r a s t to m a n y r e p o r t s s h o w i n g t h e i n t e r f e r e n c e of t r a n s p o r t by
g r a f t u n i o n s , C o l b y (1935), F r i e d r i c h (1939) a n d H e r r e r o (1951) h a v e d e m ­
o n s t r a t e d l i t t l e o r no o b s t r u c t i o n by t h e g r a f t u n io n u n d e r t h e i r c o n d i t i o n s .
T h e r e w a s no m e a s u r a b l e o b s t r u c t i o n to t h e d o w n w a r d m o v e m e n t of c a r b o ­
h y d r a t e s by t h e g r a f t u n io n in C o l b y ’s w o r k .

However, upw ard m o v em en t

m a y be l i m i t e d in d w a r f t r e e s du e to e a r l i e r s u b e r i z a t i o n of t h e i r r o o t s .
T h i s i s p a r t i c u l a r l y t r u e w h e n M a i l i n g IX i s u s e d a s a r o o t s t o c k .

H errero

(1951) f a i l e d to c o r r e l a t e t h e s t a r c h c o n t e n t a b o v e o r b e l o w t h e g r a f t u n io n
w i t h s t r u c t u r a l c o n t i n u i t y o f t h e u n io n .

He f o u n d d i f f e r e n c e s o n l y a t c e r t a i n

s e a s o n s a n d t h e s e w e r e no t a p p a r e n t d u r i n g t h e r e m a i n d e r of t h e y e a r .
T h a t r i n g s of p h l o e m f r o m th e s t o c k c a n p r o d u c e a s t o c k e f f e c t h a s
b e e n s h o w n by S w a r b r i c k (1 927a ,

1927b) a n d R o b e r t s (1934,

1935).

A ls o ,

10

p h l o e m r i n g s f r o m t h e s a m e t r e e if i n v e r t e d w i l l p r o d u c e a d w a r f i n g e f f e c t
( J a n s e 1914, Sa x 1954, a n d Sa x a n d D i c k s o n 1956).

Sax a n d D i c k s o n (1956)

f e e l t h a t t h e i n v e r t e d p h l o e m c h e c k s t h e d o w n w a r d flow of o r g a n i c n u t r i e n t s .
T h e y p o s t u l a t e t h a t by v i r t u e of r e v e r s e d p o l a r i t y (du e to i n v e r t i n g th e p h l o e m )
m o v e m e n t of o r g a n i c n u t r i e n t s a n d a u x i n s w e r e b l o c k e d .

F u rth e rm o re , foliar

a p p l i e d p h o s p h o r u s w a s s h o w n to a c c u m u l a t e i m m e d i a t e l y a b o v e th e i n v e r t e d
r i n g ( D i c k s o n a n d S a m u e l s 1956).

It i s i n t e r e s t i n g to n o t e t h a t r i n g i n g a l o n e

w i l l r e s u l t in t h e a c c u m u l a t i o n of s t a r c h a n d r e d u c i n g s u g a r s a n d in t h e
s w e l l i n g of t h e s t e m a b o v e th e r i n g (Kn igh t 1803, M a c D a n i e l s a n d C u r t i s 1930,
a n d S w a r b r i c k 1927a,
f la v in ,

1927b).

T h iam in , p y rid o x in e, p a n to th e n ic acid, r i b o ­

s u c r o s e , t o t a l n i t r o g e n a n d n o n - p r o t e i n n i t r o g e n w e r e a l l fo und to

a c c u m u l a t e in t h e v i c i n i t y of the g i r d l e d s t e m in t o m a t o p l a n t s ( B o n n e r 1944).
T h i s o c c u r r e d a s a r e s u l t of th e l i m i t e d q u a n t i t y of c o n d u c t i n g t i s s u e p r e ­
s e n t a f t e r th e r i n g i n g o r g i r d l i n g o p e r a t i o n .

T h e se effects w ere m in im ize d

a s new conducting tis s u e was g e n erate d .
B iochem ical specificity:

Im m unity re a c tio n s betw een specific p r o ­

t e i n s h a v e b e e n u s e d f o r a n u m b e r of y e a r s in a n i m a l p h y s i o l o g y to d i f f e r e n ­
t i a t e b e t w e e n s p e c i e s of a n i m a l s .

G r e e n (1926) s u g g e s t e d t h a t t h i s s a m e

r e a c t i o n m a y b e e f f e c t i v e l y u s e d in g r a f t e d p l a n t s .

P l a n t s of t h e g e n u s C i t r u s ,

which i n t e r g r a f t s e asily , gave a unifo rm p o s itiv e p r e c i p i t i n r e a c tio n with an
i m m u n e s e r u m o b t a i n e d f r o m o ne of t h e m .

A l s o , s p e c i e s of t h e s u b - f a m i l y

11

P r u n o i d e a e , kno w n to i n t e r g r a f t s u c c e s s f u l l y , g a v e u n i f o r m p o s i t i v e p r e c i p i t i n
reactions.

H o w e v e r , w h e n s e r u m f r o m t h i s g r o u p w a s a d d e d to e x t r a c t s f r o m

s p e c i e s of t h e s u b - f a m i l y P o m o i d e a e n e g a t i v e r e s u l t s w e r e o b t a i n e d .

G reen

c o n c l u d e d t h a t t h e a b s e n c e of a p r e c i p i t i n b e t w e e n t h e s e tw o s u b - f a m i l i e s m a y
be a s s o c i a t e d w i t h t h e d i f f i c u l t y o b s e r v e d in i n t e r g r a f t i n g s p e c i e s of t h e s e
tw o s u b - f a m i l i e s .

G r e e n ’s e v i d e n c e w o u l d o n ly e x p l a i n i n c o m p a t i b i l i t y w h e n

p l a n t s a r e g r a f t e d o u t s i d e of t h e i r s p e c i e s , h o w e v e r , m a n y c o m b i n a t i o n s
w i t h i n t h e s a m e s p e c i e s a r e i n c o m p a t i b l e ( T u k e y a n d B r a s e 1943).
(1928,

1929,

Kostoff

1933) p o s t u l a t e d t h a t p o s i t i v e p r e c i p i t i n r e a c t i o n s m a y be i n ­

d u c e d by g ra ftin g .

P r e c i p i t i n s not n o r m a l l y d e t e c t a b l e c a n be d e t e c t e d a f t e r

g r a f t i n g a s c i o n ( g i v i n g a p o s i t i v e p r e c i p i t i n r e a c t i o n ) on a s t o c k , w h i c h
gave a negative p re c ip itin reaction.

T h i s K o s t o f f (1929) a t t r i b u t e d to p r o ­

d u c t i o n of a n t i b o d i e s w h i c h e x e r t e d a n i n f l u e n c e on t h e s c i o n .

G enerally a

g r e a t e r d e g r e e of s p e c i f i c i t y i s a s s o c i a t e d w ith t h e p r e c i p i t i n s a c q u i r e d
through grafting.

K o s t o f f ’s c o n c l u s i o n s h a v e b e e n q u e s t i o n e d a n u m b e r of

t i m e s ( C h e s t e r 1932a,

1932b, C h e s t e r a n d W h i t a k e r 1933, W h i t a k e r a n d

C h e s t e r 1933, a n d J o n e s 1934).

C h e s t e r (1932 a) a n d C h e s t e r a n d W h i t a k e r

(1933) r e p o r t e d t h a t t h e p r e c i p i t a t e is not p r o t e i n in n a t u r e , but c h i e f l y
calcium oxalate.

T h e y d id not a g r e e t h a t t h i s so c a l l e d i m m u n o l o g y c a n

be g r a f t i n d u c e d .

T h e y f u r t h e r co n clu d e that p u b lis h e d im m u n o lo g ic a l i n t e r ­

p r e t a t i o n s o f p r e c i p i t i n r e a c t i o n s in p l a n t s a r e not t e n a b l e in v i e w of th e

12

l a c k of h o m o l o g y b e t w e e n p l a n t s a n d a n i m a l s a n d s e c o n d l y of t h e l a r g e n u m b e r
of s i m p l e n o n - p r o t e i n r e a c t i o n s p r e s e n t in p l a n t s .

F u r t h e r m o r e , t h e r e i s no

i n d i c a t i o n t h a t a s a r e s u l t of g r a f t i n g t h e r e i s a n a c q u i r e d i m m u n i t y of s t o c k
to s c i o n o r s c i o n to s t o c k d e t e c t a b l e by t h e p r e s e n t p r e c i p i t i n t e c h n i q u e .
M o r e r e c e n t l y , M i l b r a i t h a n d Z e l l e r (1945) a n d D e S t i g t e r (1956)
s u g g e s t e d l a t e n t v i r u s a n d s p e c i f i c s u b s t a n c e a s f a c t o r s w h i c h m a y be a s s o ­
c ia t e d with in c o m p a tib ility .

D w a rfe d and r o s e t t e d p e a c h t r e e s r e s u l t e d when

v i r u s f r e e b u d s w e r e p l a c e d on v i r u s i n f e c t e d s t o c k s ( M i l b r a i t h a n d Z e l l e r
1945).

F o r s u c c e s s f u l g r a f t s of m u s k m e l o n on M a l a b a r g o u r d , o n e l e a f m u s t

be p r e s e n t on th e g o u r d s t o c k ( D e S t i g t e r 1956).
the sieve tubes d is in te g ra te d .

In t h e a b s e n c e of f o l i a g e

G row th and other physiological p r o c e s s e s

r e s p o n d e d d i r e c t l y to t h e n u m b e r of l e a v e s on th e s t o c k .

DeStigter concluded

t h a t t h e p r e s e n c e of s t o c k l e a v e s i s r e q u i r e d to p r o v i d e th e s t o c k w i t h s o m e
" s p e c i f i c s u b s t a n c e " , w h i c h e n a b l e d t h e p h l o e m of th e s t o c k to f u n c t i o n n o r ­
m ally.

In t h e a b s e n c e of t h i s " s p e c i f i c s u b s t a n c e " s o m e e n z y m a t i c c o m p l e x

is a l t e r e d in t h e s i e v e t u b e s w h i c h r e s u l t s in p h l o e m d i s i n t e g r a t i o n a n d
s ta r c h accum ulation.
Ph y sio lo g ical o r individuality a s p e c ts have been p ro p o s e d .
(1894,

Daniel

1923) n o t e d t h a t p l a n t s of the s a m e f a m i l y c a n g e n e r a l l y u t i l i z e t h e

s t o r a g e r e s e r v e s of t h e r o o t s t o c k ’a s i t s own.

Along th e s e s a m e lines,

V d c h t i n g (1894) i l l u s t r a t e d t h e i n d i v i d u a l i t y of t h e g r a f t c o m p o n e n t by r e c i p r o c a l

J3

g r a f t s of s u n f l o w e r a n d a r t i c h o k e .

I r r e s p e c t i v e of t h e c o m b i n a t i o n ,

w a s fo u n d in t h e s u n f l o w e r a n d i n u l i n e in t h e a r t i c h o k e .
g r a f t s of t o m a t o a n d t o b a c c o , D a w s o n (1942,

starch

W ith r e c i p r o c a l

1945) d e m o n s t r a t e d t h a t n i c o t i n e

w a s p r o d u c e d in t h e r o o t s of t h e t o b a c c o p l a n t , a n d a c c u m u l a t e d in t h e t o m a t o
scion.

T h u s , t h e s y n t h e s i s m e c h a n i s m w a s l o c a l i z e d , a n d it w a s s h o w n t h a t

th e g r a f t u n io n p e r m i t t e d p a s s a g e of t h e a l k a l o i d .
R o o t s t o c k e f f e c t s on n u t r i e n t a b s o r p t i o n a n d a c c u m u l a t i o n :

In t h e

e a r l y p a r t of t h i s c e n t u r y D a n i e l a n d T h o m a s (1902) s u g g e s t e d t h a t g r a f t i n g
m a y m o d i f y th e p h y s i o l o g y of th e n e w l y f o r m e d p l a n t so a s to a l t e r n u t r i e n t
a b s o r p tio n and subsequent utilization.

D i f f e r e n c e s in o r g a n i c a n d i n o r g a n i c

c o m p o s i t i o n of s c i o n s on v a r i o u s r o o t s t o c k s h a s b e e n i l l u s t r a t e d on a n u m b e r
of o c c a s i o n s .

M o r e o v e r , c e r t a i n r o o t s t o c k s (e. g. p e a c h w h e n u s e d a s a

r o o t s t o c k f o r a l m o n d ) h a v e b e e n s h o w n to a c c u m u l a t e s p e c i f i c i o n s to t h e
e x t e n t t h a t t h e y a r e t o x i c to th e s c i o n , t h e r e f o r e ,

m a k in g them u n d e sira b le

a s u n d e r s t o c k s ( H a n s e n 1955).
R o a c h (1931) s t u d i e d t h e e l e m e n t s a b s o r b e d f r o m t h e s o il by s p e c t o g r a p h i c a n a l y s i s of a p p l e s c i o n s ( v a r . P r i n c e A l b e r t ) on M a i l i n g IX a n d XII
rootstocks.

T w e n t y - o n e e l e m e n t s w e r e r e c o v e r e d n a m e l y : K, Ca, Mg, Na,

P, F e , Al, Mn, Ba,

S r , L i, Cu, Pb, T i , V, Si, N, C r , Sn, M o a n d Ag .

Lead

w a s r e s t r i c t e d to t h e r o o t s , m o l y b d e n u m o c c u r r e d o n ly in t h e s t o c k p o r t i o n
of P r i n c e A l b e r t / M a i l i n g IX c o m b i n a t i o n , a n d w a s t o t a l l y a b s e n t in t h e s c i o n

14

a n d s t o c k of M a i l i n g XII.

T h e p h y s i o l o g i c a l s i g n i f i c a n c e of t h e s e o b s e r v a t i o n s

have not been e s ta b lis h e d .
U s i n g p o t a s s i u m b r o m i d e to i n d i c a t e q u a n t i t a t i v e a b s o r p t i o n ,

Berry

(1938) r e p o r t e d 1. 425, 0. 999 a n d 0. 933 m i l l i g r a m e q u i v a l e n t s of b r o m i n e a b ­
s o r b e d by M a i l i n g XIII, II a n d IX r e s p e c t i v e l y .

T h u s , he s u g g e s t e d t h a t v i g o r

of r o o t s t o c k s m a y be r e l a t e d to t h e i r c a p a c i t i e s f o r a b s o r b i n g m i n e r a l s o l u t e s .
D u r i n g t h e e a r l y p a r t of t h e s e a s o n , t r e e s on M a i l i n g XII a b s o r b e d o n e - t h i r d
m o r e n i t r o g e n a n d p o t a s s i u m t h a n t h o s e on IX, but t h e g r o w t h r a t e w a s p r o ­
p o r t i o n a l , t h e r e f o r e , no d i f f e r e n c e s w e r e o b s e r v e d in c o m p o s i t i o n on a
f r e s h w e i g h t b a s i s ( P e a r s e 1939).

A n a l y s i s of t h e a s h c o n s t i t u e n t s of t h e t e r ­

m i n a l s h o o t s o f P r i n c e A l b e r t on M a i l i n g IX a n d XII r e v e a l e d h i g h e r q u a n t i ­
t i e s of l i m e a n d a l o w e r c o n t e n t o f p o t a s s i u m in t h e s c i o n s on M a i l i n g IX.
T h e c o n t e n t of p h o s p h o r i c a c i d s h o w e d a n e g a t i v e c o r r e l a t i o n a n d m a g n e s i u m
a p o s i t i v e c o r r e l a t i o n w it h v i g o r ( V a i d y a 1938).

W a r n e a n d W a l l a c e (1935)

s t u d i e d th e i n f l u e n c e of r o o t s t o c k u p o n t h e m i n e r a l c o m p o s i t i o n of t h e s c i o n .
T h e s e a u t h o r s s h o w e d fe w d i f f e r e n c e s in c o m p o s i t i o n a n d c o n c l u d e d t h a t t h e
d w a r f i n g e f f e c t of t h e r o o t s t o c k c o u l d not be e x p l a i n e d on th e b a s i s of c h e m i ­
cal com position.

In c i t r u s , no s i g n i f i c a n t d i f f e r e n c e s in m i n e r a l c o m p o s i t i o n

have been d e m o n s tra te d am o n g cuttings,
a g i v e n s p e c i e s ( W a l l a c e et_ a ^ 1952).
f r o m the soil,

s e e d l i n g s , o r s e l f b u d d e d t r e e s of

In t h e a b s o r p t i o n of r a d i o p h o s p h o r u s

F u l l e r a n d H i l g e m a n (1955) s h o w e d no d i f f e r e n c e s in n a v e l

15

o r a n g e t r e e s on s o u r o r a n g e o r l e m o n r o o t s t o c k s , n o r d i d t h e y f in d a d i f f e r ­
e n c e in l e a f n i t r o g e n .
A c c u m u l a t i o n of s p e c i f i c i o n s a s a r e s u l t of r o o t s t o c k h a s b e e n d e m o n ­
strated.

B o ro n a c c u m u l a t e d in t h e s c i o n l e a v e s of V a l e n c i a o r a n g e on C l e o p a t r a

m a d a r i n s t o c k , w h e r e a s c h l o r i d e a n d p o t a s s i u m a c c u m u l a t e d in t h e s c i o n s on
s o u r o ra n g e ro o ts to c k s (Cooper,
1954).

G o r t o n a n d O l s o n 1952, G o r t o n a n d C o o p e r

E a t o n a n d B l a i r (1935) d e m o n s t r a t e d t h a t t h e g r a f t c o m p o n e n t s d i r e c t l y

influenced boron accum ulation.

W he n s u n f l o w e r w a s g r a f t e d o n i t s own r o o t s ,

684 p a r t s p e r m i l l i o n of b o r o n w a s r e c o v e r e d in t h e l e a v e s , c o m p a r e d to 1026
p a r t s p e r m i l l i o n w h e n g r a f t e d on a r t i c h o k e .

R e c i p r o c a l g r a f t i n g r e v e a l e d 893

p a r t s p e r m i l l i o n of b o r o n f o r a r t i c h o k e on a r t i c h o k e ,
p e r m illio n for a r tic h o k e on su nflo w e r.

c o m p a r e d to 663 p a r t s

M ore recently, alm ond rootstocks

h a v e b e e n r e c o m m e n d e d f o r a l m o n d s in l o c a t i o n s w h e r e e x c e s s b o r o n i s a
problem ,

s i n c e a l m o n d on p e a c h r e s u l t e d in s e v e r e b o r o n i n j u r y ( H a n s e n 1955).

O r g a n i c c h e m i c a l a n a l y s i s h a s f a i l e d to e x p l a i n th e d w a r f i n g e f f e c t of
c e r t a i n r o o t s t o c k o t h e r t h a n t h e a c c u m u l a t i o n of s t a r c h a b o v e t h e g r a f t u n i o n
i n a few i n s t a n c e s .

Q u a n t i t a t i v e d i f f e r e n c e s e x i s t e d in r e d u c i n g s u b s t a n c e s .

T h e l e a v e s of t h e d w a r f i n g a p p l e r o o t s t o c k s h a d a c o n s i s t e n t l y h i g h e r c o n t e n t
of r e d u c i n g s u b s t a n c e s ( L e n n a r d 1938).
c y c l e in r e d u c i n g s u g a r s ,

sucrose,

S m y t h (1938) r e p o r t e d a u n i f o r m

starch, hem icellulose, r e s e r v e c a r b o ­

h y d r a te s , c e llu lo s e an d lignin fo r a ll s to c k s te sted .

Starch and h em icellulose

16

a c c u m u l a t e d , b e g i n n i n g a b o u t th e e n d of Jun e, m o r e r a p i d l y in t h e a p p l e
s c i o n s on M a i l i n g IX t h a n i n s c i o n s on M a i l i n g II, V o r VII.
w ere re p o rte d for nitrogen fractions.

Sim ilar re su lts

In t h e c a s e of M a i l i n g IX t h e r e w a s a

t e n d e n c y of s c i o n s to a c c u m u l a t e a c i d a m i d e .

H o w e v e r , K e n c h (1938) c o n ­

c lu d e d th at the total c o n c e n tr a tio n of the v a r i o u s n itro g e n c o n s titu e n ts w as
i n d e p e n d e n t of r o o t s t o c k .
R a o a n d B e r r y (1940) i n v e s t i g a t e d t h e c a r b o h y d r a t e r e l a t i o n s of a
s i n g l e a p p l e s c i o n ( v a r . C r a w l e y Beauty) on r o o t s t o c k s M a i l i n g IX a n d XIII.
H i g h e r p e r c e n t a g e s of r e d u c i n g s u g a r s , d i - s a c c h a r i d e s a n d s t a r c h w e r e
o b s e r v e d in t h e s c i o n s on M a i l i n g IX.

A c c u m u l a t i o n of s t a r c h t o o k p l a c e

e a r l i e r i n t h e s t o c k of M a i l i n g IX, a n d a l s o in t h e s c i o n g r a f t e d t h e r e o n .
T h e y p o s t u l a t e d t h a t t h e e x c e s s i v e a c c u m u l a t i o n o f s t a r c h m a y b e th e c a u s e
of g ro w th c e s s a tio n .

S t a r c h a c c u m u l a t e d a s a r e s u l t o f a low w a t e r c o n t e n t

c a u s e d by e a r l y s u b e r i z a t i o n of M a i l i n g IX r o o t s .

T h e s t o c k i s u n a b l e to

m a i n t a i n a n a d e q u a t e w a t e r s u p p l y , t h e r e f o r e , t h e s c i o n c o n t i n u e s to g r o w
a t t h e e x p e n s e of t h e w a t e r in th e s t o c k , w h i c h in t u r n p r o m o t e s s t a r c h a c ­
c um ulation a nd f u r th e r r e s t r i c t s ro o t growth.

T h e y fo u n d no e v i d e n c e t h a t

t h e g r a f t uni on , p e r se , i n t e r f e r e s w i t h t h e d o w n w a r d t r a n s p o r t of c a r b o ­
hydrates.

F r o m t h e s e r e s u l t s they s u g g e s t th a t t h e r e w a s a la c k of m e t a ­

bolic unity b e tw ee n the scion a n d stock, with w a te r deficiency a p p e a r in g
f i r s t in t h e s t o c k a n d t h e n i n t h e s c i o n .

17

T h e a b o v e r e s u l t s a p p e a r e d to t a k e o n a d d e d i m p e t u s w h e n o n e c o n ­
s i d e r s t h e w o r k o f H a r t w e l l (1916), H a r v e y (1923) a n d H o o k e r (1926), w ho
a g r e e t h a t t h e r e t a r d a t i o n o f g r o w t h , a s a r e s u l t of a n y c a u s e , w a s a c c o m p a n i e d
by a n i n c r e a s e in s t a r c h .

It h a s n e v e r b e e n d e t e r m i n e d w h e t h e r t h i s i n c r e a s e

in s t a r c h i s t h e c a u s a t i v e f a c t o r , o r t h e r e s u l t i n g f a c t o r .

On t h e o t h e r h a n d,

R a o a n d B e r r y (1940) do n o t a t t e m p t to e x p l a i n t h e d w a r f i n g e f f e c t p r o d u c e d
b y M a i l i n g IX i n t e r s t e m a l o n e , w h i c h i s i n d e p e n d e n t of t h e r o o t s t o c k u s e d .
T h e a b i l i t y of w a t e r to m o v e i n t o t h e s c i o n h a s b e e n c o n s i d e r e d a s a
p o s s i b l e s o u r c e of s t o c k i n f l u e n c e in d w a r f a p p l e t r e e s .

T ra n sp ira tio n studies

( K n i g h t 1925, H a f e k o s t 1933) s h o w e d t h a t d w a r f a n d v i g o r o u s s t o c k s h a d d i f ­
f e r e n t t r a n s p i r a t i o n r a t e s , b u t t h a t a s i n g l e s c i o n v a r i e t y g r a f t e d on d w a r f
a n d v i g o r o u s s t o c k s s h o w e d no d i f f e r e n c e s i n t r a n s p i r a t i o n .

T his indicated

t h a t w a t e r m o v e m e n t to t h e top w a s n o t a f a c t o r l i m i t i n g g r o w t h i n d w a r f
trees.

P e a r s e (1939) u s i n g t r e e s g r a f t e d on d w a r f a n d v i g o r o u s r o o t s t o c k s ,

g r o w n in w a t e r c u l t u r e s , r e p o r t e d t h a t t r e e s on v i g o r o u s r o o t s t o c k s a b s o r b e d
m o r e w a t e r p e r u n i t l e a f a r e a t h a n t h o s e on d w a r f i n g r o o t s t o c k s .

T r e e s on

d w a r f i n g r o o t s t o c k s , h o w e v e r , a b s o r b e d m o r e w a t e r p e r u n it i n c r e a s e in
f r e s h w eight th a n th o s e on v ig o ro u s ro o ts to c k s .

18

T ra n s lo c a tio n Studies

G e n e r a l c o n s i d e r a t i o n s on t r a n s p o r t :

S i n c e t h e c l a s s i c a l s t u d y of

H e v e s y (1923) s h o w i n g t h e a b s o r p t i o n a n d s u b s e q u e n t t r a n s p o r t of t h o r i u m B
by p l a n t s , i n c r e a s i n g e m p h a s i s h a s b e e n g i v e n to t r a n s p o r t s t u d i e s in b i o l o ­
gical sy ste m s .

T h is in d ic a te s that th e se tr a n s p o r t p r o c e s s e s a r e equally

i m p o r t a n t to t h e m u l t i t u d e of r e a c t i o n s t a k i n g p l a c e in v a r i o u s p l a n t t i s s u e s .
T r a n s p o r t s t u d i e s , in r e l a t i o n to s t o c k - s c i o n e f f e c t , w e r e g i v e n a t r e m e n d o u s
i m p e t u s w h e n S w a r b r i c k a n d R o b e r t s (1927) d e m o n s t r a t e d t h a t a d w a r f i n g
i n t e r s t e m p l a c e d b e t w e e n a v i g o r o u s s c i o n a n d s t o c k w o u l d d w a r f th e s c i o n .
L a t e r , R o b e r t s (1934) d e m o n s t r a t e d t h a t a r i n g of b a r k f r o m th e s t o c k c o u l d
also p ro d u c e a dw arfing effect.

T h e p h y s i o l o g i c a l s i g n i f i c a n c e of t h e s e f i n d ­

i n g s w e r e no t a s c e r t a i n e d .
E a r l y in t h e e i g h t e e n t h c e n t u r y H a l e s (1727) r e p o r t e d t h a t w a t e r m o v e d
u p w a rd th ro u g h the wood and do w n w ard th ro u g h the b a rk .

K n ig h t (1803,

c o n f i r m e d t h e s e r e s u l t s a n d a t t e m p t e d to a c c o u n t f o r s a p m o v e m e n t .

1830)

This

l o n g s t a n d i n g b e l i e f w a s q u e s t i o n e d b y C u r t i s (1920), Z i m m e r m a n a n d C o n n a r J
(1934), G u s t a f s o n a n d D a r k i n (1937), a n d G u s t a f s o n (1939).

G ustafson and

D a r k i n (1937) w e r e c o n v i n c e d t h a t p h o s p h o r u s p a s s e d u p w a r d t h r o u g h th e
phloem a s well a s the xylem .

T h e y m a i n t a i n e d t h a t th e p h l o e m a n d x y l e m

f u n c t i o n e d in t h e u p w a r d m o v e m e n t of p h o s p h o r u s in w il lo w a n d g e r a n i u m .
Z i m m e r m a n a n d C o n n a r d (1934), u s i n g i n a r c h e d t o m a t o a n d c o l e u s p l a n t s ,

19

c o n c lu d e d that e l a b o r a t e d foods a s well a s m i n e r a l s a lts m o v e d u p w a rd and
downw ard through s te m s.

T h e o r i e n t a t i o n of t h e i n t e r v e i n i n g s t e m w a s c o n ­

s i d e r e d of no i m p o r t a n c e ,

sin ce s o m e a c tiv ity at a d is ta n t point w a s the

d e t e r m i n i n g f a c t o r of t r a n s p o r t .

D ix o n (1922) a n d Dix on a n d Ball (1922)

d e m o n s t r a t e d t h a t t h e x y l e m f u n c t i o n e d in d o w n w a r d t r a n s p o r t of o r g a n i c
m aterials.

E o s i n d ye a p p l i e d to t h e c u t tip of p o t a t o a n d c h r y s a n t h e m u m

a p p e a r e d t h r o u g h o u t t h e p l a n t w i t h i n 24 h o u r s , a n d w a s fo und in th e v e s s e l s .
T h e y c o n c l u d e d t h a t r e v e r s e t r a n s p i r a t i o n r e s u l t e d in t h e d o w n w a r d m o v e ­
m e n t of w a t e r in t h e v e s s e l s , a n d t h i s e f f e c t e d th e d o w n w a r d t r a n s p o r t of
solutes.

T h e c l a s s i c a l w o r k of St out a n d H o a g l a n d (1939) u s i n g r a d i o ­

i s o t o p e s of P, Na, Br a n d K, i l l u s t r a t e d c l e a r l y t h a t th e x y l e m w a s th e
p r i m a r y u p w a r d p a t h w a y f o r t r a n s p o r t in a c t i v e l y g r o w i n g p l a n t s whe n t h e
b a r k and xylem w e re se p a ra te d .

H o w e v e r , if th e b a r k w a s le ft i n t a c t ,

l a t e r a l m o v e m e n t r a p i d l y o c c u r r e d , a n d t h e i s o t o p e w a s p r e s e n t in both
xylem and phloem .
C

14

C h e n (1951) s h o w e d s i m u l t a n e o u s m o v e m e n t of P

in t h e p h l o e m of w i l l o w a n d g e r a n i u m c u t t i n g s .

oo

and

M o r e r e c e n t l y , both

b a s i p e t a l a n d a c r o p e t a l t r a n s p o r t of p h o s p h o r u s in th e p h l o e m of c o t t o n w a s
d e m o n s t r a t e d by Biddulph a n d M a r k l e (1944).

A u t o r a d i o g r a m s p r e p a r e d of

t h e b e a n s t e m s j u s t b e l o w t h e n o d e of S 35 a n d P 32 t r e a t e d l e a v e s , i n d i c a t e d
t h a t t h e f i r s t p a t h w a y of d o w n w a r d m o v e m e n t of s u l f u r a n d p h o s p h o r u s w a s
th ro u g h the p h lo e m .

T h e r e w a s t h e n d i f f u s i o n in to t h e x y l e m (Biddulph 1956,

a n d Biddulph, C o r y a n d Bid dul ph 1956).

20

T r a n s p o r t of p h o s p h o r u s : H e v e s y et a l (1936) s u g g e s t e d t h a t th e
p h o s p h o r u s a t o m s in p l a n t s a r e in a c o n t i n u o u s i n t e r c h a n g e b e t w e e n th e d i f ­
fe re n t p la n t p a r t s , thus concluding that p h o s p h o ru s is a v e r y m obile e lem en t.
T h e s e f i n d i n g s h a v e b e e n c o n f i r m e d by a l a r g e n u m b e r of w o r k e r s ( A r n o n et al
1940, C o l w e l l 1942, S i l b e r s t e i n a n d W i t t w e r 1951, N o r t o n 1954 a n d Biddulph
1955).

B id du lp h (1941) s h o w e d a d i u r n a l m i g r a t i o n of

l e a v e s of b e a n p l a n t s .

i n j e c t e d in to t h e

T h e g r e a t e s t d o w n w a r d m i g r a t i o n w a s a t o r n e a r 10:0 0

A . M . w it h th e l e a s t a t o r n e a r 10:00 P . M .

F u r th e r m o r e , ph o sp h o ru s applied

to t h e f o l i a g e of p l a n t s i s r e a d i l y a b s o r b e d a n d u t i l i z e d in t h e m e t a b o l i c p r o ­
c e s s e s of t h e p l a n t .
of p lan ts,

T h i s h a s l e d to t h e u t i l i z a t i o n of s u p p l e m e n t a l f e e d i n g

by f o l i a r a p p l i c a t i o n s , d u r i n g c r i t i c a l s t a g e s in t h e i r o n t o g e n y

(S ilb e rstein and W ittwer,
1956).

1951, E g g e r t et a l 1952, A s e n e t al 1953, K u r s a n o v

T e m p e r a t u r e ( S w a n s o n a n d W h i t n e y 1953), t r a n s p i r a t i o n ( W r i g h t a n d

B a r t o n 1955), m e t a b o l i c i n h i b i t o r s ( K e n d a l l 1955), a n d g e n e r a l h e a l t h (S t a n k o v i c
a n d B e c a r e v i c 1954) of t h e p l a n t a r e a l l f a c t o r s w h i c h i n f l u e n c e t h e a b s o r p t i o n ,
t r a n s p o r t a n d d i s t r i b u t i o n of p h o s p h o r u s .

T h e s e e f f e c t s m u s t be c o n s i d e r e d

in t h e i n t e r p r e t a t i o n of r e s u l t s .
T r a n s p o r t of c a l c i u m :
p la n ts , c a lc iu m is im m o b ile .
calcium o ccu rs.

In c o n t r a s t to p h o s p h o r u s w h i c h i s m o b i l e in
V e ry little,

if any, d o w n w a r d t r a n s p o r t of

T h e n e c e s s i t y of c a l c i u m in t h e r o o t m e d i a w a s d e m o n s t r a t e d

by H a y n e s a n d R o b i n s (1948), R i r i e a n d T o t h (1952) a n d B l e d s o e (1953).

21

G r o w i n g h a l f of t h e r o o t s y s t e m of a s i n g l e t o m a t o p l a n t in a c o m p l e t e
n u t r i e n t s o l u t i o n a n d t h e o t h e r h a l f in a n u t r i e n t s o l u t i o n m i n u s c a l c i u m ,
R i r i e a n d T o t h d e m o n s t r a t e d t h e i n a b i l i t y of c a l c i u m to m o v e in to t h e r o o t s
g r o w i n g in t h e c a l c i u m d e f i c i e n t s o l u t i o n .
a b s o r p t i o n o f c a l c i u m by th e p e a n u t f r u i t ,

T h e s e r o o t s d ie d.

S t u d y i n g th e

B le d s o e e t a l (1949) c o n c l u d e d

t h a t t h e f r u i t i n g p e g w a s d e p e n d e n t on th e p e g m e d i u m f o r i t s c a l c i u m
supply.
Bean p l a n t s g r o w n in a n u t r i e n t s o l u t i o n c o n t a i n i n g C a
g r o w n in a s o l u t i o n c o n t a i n i n g n o n - r a d i o a c t i v e c a l c i u m ,
c i a b l e m o v e m e n t of C a

45

45

a n d then

s h o w e d no a p p r e -

i n to th e n e w l y d e v e l o p e d t i s s u e .

C alcium a sc e n d s

t h r o u g h th e x y l e m to th e l e a v e s , but t h e r e i s no e v i d e n c e t h a t it i s r e - e x p o r t e d
f r o m th e l e a f v i a t h e p h l o e m ( M a s o n a n d M a s k e l l 1931).

In fa c t, t h e s e

a u t h o r s c o n c l u d e d t h a t th e i m m o b i l i t y of c a l c i u m w i t h i n the p l a n t is a s s o c i a t e d
w i t h i t s i n a b i l i t y to m o v e in t h e p h l o e m .

O n c e c a l c i u m i s in t h e l e a f it is

e i t h e r p r e c i p i t a t e d o r c o m b i n e d w i t h t i s s u e m a t e r i a l so t h a t few f r e e c a l ­
c i u m i o n s a r e le f t in s o l u t i o n .

S m i t h (1944) s u g g e s t e d t h a t b o r o n i s a s s o ­

c i a t e d w i t h t h e s o l u b i l i t y o f c a l c i u m p a r t i c u l a r l y in th e c y t o p l a s m a n d c e l l
wall.

H o w e v e r , W i t t w e r (1956) f o u n d t h a t b o r o n d id not f a c i l i t a t e t h e b a s i -

p e t a l m o v e m e n t of c a l c i u m in b e a n p l a n t s .

T h e r e is c o n s i d e r a b l e e v id en c e

t h a t t h e t r a n s p o r t of o r g a n i c c o m p o u n d s in p l a n t s m a y be r e g u l a t e d by b o r o n
( G a u c h a n d D u g g e r 1954, S i s l e r _ e t j i l 1 956), h o w e v e r , t h e m e c h a n i s m r e ­
m ains obscure.

22

T ran sp o rt and p o larity :

P o l a r t r a n s p o r t of n a t u r a l l y o c c u r r i n g

a u x i n s in p l a n t s h a s b e e n k n o w n f o r a n u m b e r of y e a r s (Wen t 1932, C l a r k
1937, a n d L e o p o l d a n d G u e r n s e y 1953).

F u rth e rm o re , th ere is a re p o rt

t h a t p o l a r i t y c a n be t e m p o r a r i l y s u s p e n d e d by m i l d a n t h e s i a ( W e nt a n d
T h i m a n n 1937), w h i c h r e s u l t s in t h e n o n - p o l a r m o v e m e n t o f a u x i n .
N i e d e r g a n g - K a m i e n a n d S koog (1956) r e p o r t e d a m o d i f i c a t i o n of a u x i n
t r a n s p o r t w hen to b a cc o s te m c y l i n d e r s a r e p r e t r e a t e d with tr i io d o b e n z o ic
acid.

No a t t e m p t h a s b e e n m a d e to e x p l a i n t h e f a i l u r e of c e r t a i n i o n s to

m o v e on t h e b a s i s o f p o l a r i t y .

H o w e v e r , M a s o n a n d M a s k e l l (1931) l e a d

o n e to s u s p e c t t h a t c a l c i u m m a y b e h a v e in s u c h a m a n n e r w h e n th e y
c o n c l u d e d t h a t t h e i m m o b i l i t y of c a l c i u m in p l a n t s i s a s s o c i a t e d w i t h
i t s i n a b i l i t y to m o v e in t h e p h l o e m .

W e n t (1932) a t t a c h e d s i g n i f i c a n c e

to p o l a r t r a n s p o r t w h e n he s t u d i e d t h e m o v e m e n t of o r g a n i c d y e s in
plants.

A cid d y e s m o v e d m o r e r a p id ly to w a r d the b a s e than to w a rd

t h e a p e x , w h e r e a s b a s i c d y e s m o v e d m o r e r a p i d l y to t h e a p e x t h a n t h e
base.

T h i s s u g g e s t e d t h a t d i r e c t i o n of m o v e m e n t i s d e t e r m i n e d not

m e r e l y by c h e m i c a l c o n s t i t u t i o n bu t a l s o by t h e n a t u r e of t h e d i s s o c i a t i n g
groups.

23

III. G E N E R A L M E T H O D S A N D M A T E R I A L S

P r e p a r a t i o n of a p p l e s t i o n s :

S t i o n s ( c o m b i n a t i o n of s t o c k a n d s c i o n ,

T u k e y 1937) w e r e p r e p a r e d by " T ” b u d d i n g c l o n a l M c I n t o s h b u d s on y e a r o l d E a s t M a i l i n g (EM) VII, IX a n d XVI a n d D e l i c i o u s s e e d l i n g r o o t s t o c k s .
T h e s t i o n s w e r e l i f t e d f r o m t h e n u r s e r y r o w in N o v e m b e r a n d p l a c e d in
a n u r s e r y s t o r a g e (40-5CPF) until used.
Bu dded a n d g r a f t e d t o m a t o p l a n t s ( v a r .

Stokesdale and M oreton

H y b r i d ) w e r e p r e p a r e d by b u d d i n g a s h i e l d c o n t a i n i n g o n e bu d to t o m a t o
s t o c k s of t h e s a m e v a r i e t y .

T h e g r a f t e d p l a n t s w e r e p r e p a r e d by g r a f t ­

i n g a t w o - i n c h s c i o n on s t o c k s of th e s a m e v a r i e t y , u s i n g th e s a d d l e o r
whip a n d t o n g u e m e t h o d (J on e s 1934).

T h e n e w l y p l a c e d bu d o r s c i o n

w a s s e c u r e l y w r a p p e d in p l a c e w i t h a l a t e x

1

b a nd .

W a te r was withheld

f r o m h e r b a c e o u s p l a n t s u s e d a s s t o c k s f o r 24 h o u r s b e f o r e g r a f t i n g .
T h i s f a c i l i t a t e d th e g r a f t i n g o p e r a t i o n by p e r m i t t i n g c l o s e r c o n t a c t of th e
c a m b i u m a r e a s of t h e s t o c k a n d s c i o n , a n d r e s u l t e d in s t r o n g e r u n i o n s .
G row ing plants:

A p p l e s t i o n s w e r e r e m o v e d f r o m th e n u r s e r y

s t o r a g e c e l l a r in F e b r u a r y .

T h e s t o c k a b o v e th e n e w l y p l a c e d b u d w a s

r e m o v e d a n d t h e s t i o n s w e r e g r o w n in s o l u t i o n c u l t u r e s .

A m odified

H o a g l a n d s o l u t i o n ( H o a g l a n d a n d A r n o n 1950) w i t h f e r r i c e t h y l e n e d i a m i n e
t e t r a a c e t a t e a s t h e i r o n s o u r c e w a s u s e d a s s u g g e s t e d by J a c o b s o n (1951).

^ M a n u f a c t u r e d by t h e S e a l t e x C o m p a n y , C h i c a g o , I l l i n o i s .

24

P l a n t s w e r e s t a r t e d in a h a l f s t r e n g t h H o a g l a n d s o l u t i o n , a n d a s s o o n a s
t h e b u d s b r o k e t h e s o l u t i o n w a s c h a n g e d to a m o d i f i e d (full s t r e n g t h )
Hoagland, a n d r e p l a c e d w eekly t h e r e a f t e r .

W h e n p l a n t s w e r e g r o w n in

s o i l c u l t u r e s h ig h n u t r i e n t l e v e l s w e r e m a i n t a i n e d by f r e q u e n t f e r t i l i ­
zation with d ia m m o n iu m a n d m o n o p o ta s s iu m p h o s p h a te s .
Isotope s tu d ie s :

Radioactive m a te ria ls ,

both p r o c e s s e d a n d i r r a ­

d i a t e d u n i t s , w e r e o b t a i n e d e i t h e r f r o m t h e O ak R i d g e N a t i o n a l L a b o r a t o r y ,
O ak Ridge, T e n n e s s e e , N u c le a r Science a n d E n g i n e e r in g C o rp o ra tio n ,
P i t t s b u r g , P e n n s y l v a n i a , o r A b b o tt L a b o r a t o r i e s ,
The P
Ca

45

32

C h i c a g o 36, I l l i n o i s .

w a s r e c e i v e d a s o r t h o - p h o s p h o r i c a c i d (H^P

C I 2 in w e a k h y d r o c h l o r i c a c i d .

“7
As
O^) a n d C a ^ 3 a s

O th e r is o to p e s w e re o b tain ed a s the

c h l o r i d e in w e a k h y d r o c h l o r i c a c i d w it h th e e x c e p t i o n of Cu

, Cl

and

S3 5 , w h i c h w e r e r e c e i v e d a s C u ^ N O g , H C l 3^ a n d S 3 3 C>4 r e s p e c t i v e l y .
In s t u d i e s i n v o l v i n g d o w n w a r d t r a n s p o r t , a d e f i n i t e q u a n t i t y of t h e
i s o t o p i c s o l u t i o n w a s a p p l i e d to t h e m i d v e i n of a le a f .

A tuberculin syringe

w i t h a n u m b e r 27 s t a i n l e s s s t e e l n e e d l e o r a m i c r o p i p e t t e w a s u s e d .

For

r o o t a b s o r p t i o n s t u d i e s , a d e f i n i t e q u a n t i t y of th e i s o t o p i c a l l y l a b e l e d
s o l u t i o n w a s a p p l i e d to t h e r o o t m e d i a .

R e s u lts from a p r e l i m i n a r y study

w e r e u s e d to e s t a b l i s h s u i t a b l e i s o t o p e a n d c a r r i e r c o n c e n t r a t i o n s a n d
t i m e i n t e r v a l b e t w e e n t r e a t m e n t a n d h a r v e s t of s p e c i f i c p l a n t t i s s u e s .
Plants t r e a t e d as d e s c rib e d above w e re h a rv e s te d at v a rio u s tim e

25

intervals after treatm ent,

s e p a r a t e d into t h e d e s i r e d s a m p l e s , a n d d r i e d

to a c o n s t a n t w e i g h t in a f o r c e d a i r o v e n a t 70® C.

Dry weights w e re d e ­

t e r m i n e d , t h e s a m p l e s c u t in to s m a l l s e g m e n t s , a n d a s h e d by s t a n d a r d d r y
o r wet p ro c e d u re s .
R a d i o c a l c i u m d e t e r m i n a t i o n s w e r e m a d e by a s h i n g t h e p l a n t s a m p l e s
in 50 m i l l i l i t e r c r u c i b l e s a t 550® C f o r 12 h o u r s in a m u f f l e f u r n a c e .
l o s s of r a d i o a c t i v i t y w a s d e t e c t a b l e u n d e r t h e s e c o n d i t i o n s .

No

The resulting

a s h w a s d i s s o l v e d in 2N h y d r o c h l o r i c a c i d , t r a n s f e r r e d to a 25 m i l l i l i t e r
v o l u m e t r i c f l a s k a n d b r o u g h t up to v o l u m e .

One o r fi v e m i l l i l i t e r a l i q u o i t s

w e r e t r a n s f e r r e d to C o o r m i l k a s h i n g c a p s u l e s * , a n d e v a p o r a t e d to a
u n i f o r m f i l m u n d e r a b a n k of 250 w a tt i n f r a r e d h e a t l a m p s .

All s a m p l e s

w e r e c o u n ted with a m ic a en d -w in d o w G e ig e r -M illle r tube u s in g a T r a c e r l a b
Model S C -IB A u to s c a le r o r N u c lea r In stru m e n t and C h em ical C o rp o ratio n ,
M o d e l 172 U l t r a s c a l e r .

Both i n s t r u m e n t s w e r e e q u i p p e d w i t h s h i e l d e d

counting c h a m b e rs .
R a d i o p h o s p h o r u s d e t e r m i n a t i o n s r e q u i r e d p r e - d i g e s t i o n of p l a n t
t i s s u e a n d f i x a t i o n of t h e p h o s p h o r u s p r i o r to a s h i n g .

Sam ples w ere tre a te d

a c c o r d i n g to t h e p r o c e d u r e d e s c r i b e d by A. O. A. C. (1950); f i r s t , 5 N m a g ­
n e s i u m n i t r a t e t h e n c o n c e n t r a t e d h y d r o c h l o r i c a c i d w e r e a d d e d to t h e s a m ­
p l e s while on a s te a m bath.

The concentrated hydrochloric acid serv ed

to p r e - d i g e s t t h e t i s s u e p e r m i t t i n g c l o s e r c o n t a c t b e t w e e n th e t i s s u e
^ C o o r m i l k a s h i n g c a p s u l e s , m a n u f a c t u r e d by C o o r ' s a n d C o m p a n y ,
Golden, C o lo ra d o .

26

p h o s p h o r u s and the a dded m a g n e s iu m n itr a te .

The m agnesium n itrate

f o r m e d m a g n e s i u m p y r o p h o s p h a t e w i t h t h e p h o s p h o r u s in t h e p l a n t t i s s u e ,
w h i c h i s t h e r m a l l y s t a b l e a t 550° C, t h u s p e r m i t t i n g c o m p l e t e a s h i n g in
a muffle fu rn ac e.

T h e a s h w a s d i s s o l v e d in 2 N h y d r o c h l o r i c a c i d a n d

t r e a t e d a s d e s c r ib e d for c alciu m above.
It w a s f o u n d m o r e c o n v e n i e n t , in w o r k i n g w i t h w oo d y m a t e r i a l ,
to w e t a s h t h e s a m p l e s .

Stem s e g m e n t s a n d g ra ft union s e g m e n ts w e r e

c u t in to s m a l l p i e c e s a n d t r e a t e d w i t h fiv e m i l l i l i t e r s of c o n c e n t r a t e d
n i t r i c a c i d p e r g r a m of t i s s u e .

A fte r the in itial r e a c t io n sub sid ed , an

a d d i t i o n a l f iv e m i l l i l i t e r s w e r e a d d e d a n d h e a t e d on a s t e a m b a t h .

The

t e m p e r a t u r e w a s g r a d u a l l y i n c r e a s e d u n t i l a l l of t h e p l a n t t i s s u e w a s o x i d i z e d
a n d the so lu tio n h a d tak en on a light yellow c o lo r (a p p r o x im a te ly

12

hours).

A f t e r c o o l i n g , t h e s a m p l e s w e r e t r a n s f e r r e d t o a 25 m i l l i l i t e r v o l u m e t r i c
fla sk and tr e a te d a s d e s c rib e d p re v io u s ly for Ca^^.
M o s t s a m p l e s w e r e c o u n t e d u n t i l 4096 o r 2048 c o u n t s h a d r e g i s t e r e d
on t h e c o u n t e r .

H o w e v e r , in c a s e of low r a d i o a c t i v i t y e a c h s a m p l e w a s

c o u n t e d f o r t h r e e m i n u t e s , o r u n t i l 400 c o u n t s h a d r e g i s t e r e d .

Counts

tw ic e the e x is tin g b a c k g ro u n d w e r e d e e m e d n e c e s s a r y for significance.
R a d io a c tiv e a n a ly s is w e r e e x p r e s s e d a s total counts p e r m in u te (cpm) o r
c o unts p e r m in u te p e r g r a m ( c p m /g m ) d ry weight.
C alculations w e re m a d e fo r ra d io a ctiv e decay when n e c e s s a ry .

27

T h e follow ing equation w as used:
At = A 0 e _>lt
W here:
A
A0
e
✓\
t

=
=
=
-

a c t i v i t y at t i m e i n t e r v a l t
activity at initial tim e
b a s e of n a t u r a l log. (2. 3026)
d e c a y c o n s t a n t f o r th e i s o t o p e u s e d
tim e elapsed

No c o r r e c t i o n s w e r e m a d e f o r s e l f - a b s o r p t i o n s i n c e a l l but two e x p e r i m e n t s
w e r e c o u n ted a s an infinitely thin s a m p le la y e r.

D i r e c t c o u n t i n g of d r i e d

p l a n t m a t e r i a l w a s s u c c e s s f u l l y e m p l o y e d in two e x p e r i m e n t s a f t e r no d i f f e r ­
e n c e s c o u l d be o b s e r v e d on a c o m p a r a t i v e b a s i s .

S e lf - a b s o r p tio n was only a

f a c t o r w h e r e p l a n t s p e c i m e n s d i f f e r e d g r e a t l y in w ei g h t.
E x p e r im e n ts w e re c o m p le tely ra n d o m iz ed , r a n d o m iz e d block o r split
plot d e sig n s.

E s t i m a t e s of v a r i a b i l i t y w e r e by th e a n a l y s i s of v a r i a n c e m e t h o d

d e s c r i b e d by S n e d e c o r (1948).

W h ere significant " F " values o c c u r re d , c a l ­

c u l a t i o n s w e r e m a d e f o r l e a s t d i f f e r e n c e s n e c e s s a r y f o r s i g n i f i c a n c e a t th e
p r o b a b i l i t y of fi v e a n d o n e p e r c e n t l e v e l s ( d e s i g n a t e d L. S. D. 5% a n d 1%
respectively).

W h e r e ” F ” v a l u e s w e r e i n s i g n i f i c a n t , th e d e s i g n a t i o n " n o t

significant" was used.
A u t o r a d i o g r a p h y w a s u s e d to s h o w g r o s s a b s o r p t i o n a n d s u b s e q u e n t
t r a n s p o r t of r a d i o i s o t o p e s f r o m th e s i t e of t r e a t m e n t ( W i t t w e r a n d L u n d a h l
1951 a n d Boyd 1955).

S a t i s f a c t o r y a u t o r a d i o g r a m s w e r e o b t a i n e d of w ood y

p l a n t m a t e r i a l by r a p i d l y d r y i n g at 80° C, c u t t i n g in to fiv e m i l l i m e t e r

28

l o n g i t u d i n a l s e c t i o n s , a n d p l a n e i n g to o b t a i n a s m o o t h s u r f a c e .

If a l l

o p e r a t i o n s w e r e m a d e in t h e d i r e c t i o n of a n i n c r e a s i n g a c t i v i t y g r a d i e n t ,
l i t t l e o r no c o n t a m i n a t i o n o c c u r r e d .

T h e s p e c i m e n s w e r e fixed on a

m o u n t i n g b o a r d , c o v e r e d w it h S a r a n w r a p (Dow C h e m i c a l C o m p a n y ,
M i d l a n d , M i c h i g a n ) a n d p l a c e d in c o n t a c t w it h X - r a y f i l m f o r e x p o s u r e .
H e rb a c e o u s s p e c im e n s w e re p r e p a r e d for a u to ra d io g ra p h y , a s d e s c r ib e d
by W i t t w e r a n d L u n d a h l (1951).
K o d a k X - r a y n o - s c r e e n o r b l u e - b r a n d fi lm w a s u s e d , d e p e n d i n g
on the s e n s itiv ity d e s i r e d .

E x p o s u r e t i m e w a s d e t e r m i n e d by p r e l i m i n ­

a r y s t u d i e s c o n s i d e r i n g t h e i s o t o p e in q u e s t i o n , a n d r a d i a t i o n i n t e n s i t y
of t h e m o u n t e d s p e c i m e n .

T h e e x p o s e d fi l m w a s d e v e l o p e d u s i n g K o d a k

X - r a y d e v e l o p e r a n d fixed with Kodak X - r a y f ix e r and r e p l e n i s h e r .

29

IV. E X P E R IM E N T A L R E SU L T S

S e c tio n A
E ffe c t o f S to c k -S c io n I n te r r e la tio n s h ip U pon N u trie n t
A b s o rp tio n a n d T r a n s p o r t

E x p e rim e n t

1

R e la tiv e a b s o rp tio n a n d t r a n s p o r t c a p a c itie s o f ro o te d c u ttin g s of
M a ilin g VII, IX, a n d XVI w e r e stu d ie d .

N ine u n ifo rm r o o te d c u ttin g s of

E a s t M a ilin g (EM ) VII, IX, a n d XVI w e re s e le c te d fro m th e n u r s e r y s t o r ­
a g e on M a rc h 24, 1955.

T h e y w e re th o ro u g h ly w ash ed , t r a n s f e r r e d to s o lu ­

tio n c u ltu r e s (6 l i t e r g la z e d c r o c k s ) an d c a r e d f o r a s d e s c r ib e d u n d e r
m e th o d s .

T h r e e d a y s a f t e r th e in itia tio n of g ro w th a ll but one of th e new

sh o o ts w e re re m o v e d .

On A p ril 27, 0. 5 m ic r o c u r ie (uc) of P 32 p e r m i l l i ­

g r a m o f P ^ l w as a d d e d to th e so lu tio n c u ltu r e ^ (a p p ro x im a te ly 93. 0 u c /
s o lu tio n c u ltu r e ) .

T h is w as b elo w th e ra n g e w h ich r e s u l t e d in r a d ia tio n

in ju ry a c c o r d in g to Blum e e t a l (1950) a n d Blum e (1952).

T h e new sh o o t

a n d r o o ts w e r e h a r v e s te d 24, 48 a n d 96 h o u rs a f te r tr e a tm e n t.

The h a r­

v e s te d s h o o ts (to p s) w e r e d rie d , w eig h ed , w et a s h e d an d a s s a y e d f o r
r a d io a c tiv ity .

D ry w e ig h ts o f th e r o o ts an d to p s w e r e r e c o r d e d .

The

r a d io a c tiv ity w a s e x p r e s s e d a s c o u n ts p e r m in u te p e r g ra m d ry w eig h t
o f to p p e r m illig r a m of r o o t.
in q u a n tity o f r o o ts .

T h is c o n s titu te d a c o r r e c t io n fo r d iffe re n c e

F ig u r e 1

A c c u m u l a t i o n of r a d i o p h o s p h o r u s (P

oo

) in t h e t o p s of

r o o t e d c u t t i n g s of E a s t M a i l i n g (EM) VII, IX a n d XVI, 24,
48 a n d 96 h o u r s a f t e r a p p l y i n g P

oo

to th e r o o t m e d i u m .

HOURS

FOLLOWING

TREATMENT

31

T h e d a t a s h o w ( F i g u r e 1) t h a t t h e t o p s of E M VII, IX a n d XVI a c c u m u l a t e d i n c r e a s i n g q u a n t i t i e s of t h e r o o t a p p l i e d P
q u a n t i t y of r o o t a p p l i e d

oo

oo

with tim e .

The g re a te s t

w a s a b s o r b e d a n d a c c u m u l a t e d by t h e v i g o r o u s

r o o t s t o c k E M XVI, a n d t h e l e a s t by t h e d w a r f i n g r o o t s t o c k E M IX.
s e m i - d w a r f r o o t s t o c k E M VII w a s i n t e r m e d i a t e .

The

No e x p l a n a t i o n c a n be

o f f e r e d a s to why the t o p s of E M VII c o n t a i n e d l e s s r a d i o a c t i v i t y a t 24 h o u r s
t h a n t h e m o r e d w a r f i n g E M IX.

E x p e r i m e n t s 2, 3 a n d 4

T h e f o l l o w i n g e x p e r i m e n t s w e r e d e s i g n e d to o b s e r v e t h e f o l i a r a b s o rp tio n and t r a n s p o r t of P
Ca

45

32

a n d r o o t a b s o r p t i o n a n d t r a n s p o r t of P

32

and

by a p p l e s c i o n s ( v a r . M c I n t o s h ) b u d d e d on d w a r f i n g (E M IX a n d VII)

a n d v i g o r o u s ( E M XVI a n d D e l i c i o u s s e e d l i n g ) r o o t s t o c k s .
T h i r t y - s i x ( n in e of e a c h r o o t s t o c k ) u n i f o r m t r e e s , w h i c h w e r e b u d d e d
in A u g u s t , w e r e r e m o v e d f r o m t h e n u r s e r y s t o r a g e c e l l a r a n d th e s t o c k
above the newly p l a c e d bud w as re m o v e d .
and uniform ly pruned.

The ro o ts w ere thoroughly w ashed

T h e y w e r e p l a c e d in 3. 5 l i t e r g l a s s j a r s c o n t a i n i n g

a h a l f - s tr e n g t h H oagland solution.

T h e j a r s c o n t a i n i n g th e p l a n t s p e c i m e n s

w e r e t h e n s e t in c o n t r o l l e d t e m p e r a t u r e t a n k s m a i n t a i n e d a t 55° F t 2. 0° by
a c i r c u l a t i n g w a t e r bath and t h e r m o s t a t i c a l l y c o n tr o lle d r e f r i g e r a t o r and
h e a t i n g c o i l s ( F i g u r e 2).

T h e d e s i g n o f th e c o n t r o l l e d t e m p e r a t u r e t a n k s

F ig u re 2

So lu ti on c u l t u r e s u s e d in g r o w i n g th e p l a n t s .

Root

m e d i u m m a i n t a i n e d at 55° F in e a c h t a n k by r e f r i g e r a t o r c o m p r e s ­
s o r s and heating units.

A Brown a u t o m a t i c p o t e n t i o m e t e r c o n t i n u ­

o u s l y r e c o r d e d t e m p e r a t u r e of w a t e r - b a t h , n u t r i e n t s o l u t i o n a n d
air.

33

h a s b e e n d e s c r ib e d in d e ta il by R o b e r ts (1953).

T e m p e r a tu r e o f th e w a te r -

bath , n u tr ie n t s o lu tio n a n d a i r w e r e r e c o r d e d a u to m a tic a lly by a Brown
P o te n tio m e te r* .

T h e n u tr ie n t s o lu tio n s w e r e v ig o ro u s ly a e r a t e d fro m a

c e n t r a l a i r su p p ly w h ich w as f il t e r e d a n d d is tr ib u te d to e a c h j a r th ro u g h
a e r a t i n g s to n e s ^ .
W ithin a w e ek a f t e r p la c in g in s o lu tio n c u ltu r e s , a fu n g u s w as o b ­
s e r v e d on th e r o o ts of EM XVI a n d D e lic io u s s e e d lin g .

It a p p e a r e d to be

lo c a liz e d on th e o ld r o o ts ; new r o o ts w e re not a ffe c te d .

T h e fu n g u s w as

is o la te d a n d id e n tifie d a s a M u c o r s t r a i n s a p ro p h y te .

A fte r re m o v in g th e

s tio n s , th e r o o ts w e r e w a s h e d a n d th e n u trie n t s o lu tio n s w e re r e p la c e d w ith
s ta n d a r d H o ag lan d .

T h e fungus w as not a s e r io u s p ro b le m once th e b u d s

b ro k e an d th e s tio n s w e re a c tiv e ly g ro w in g .

A lso , in a n u m b e r of c a s e s th e

new ly p la c e d b u d a p p e a r e d a s a m ix e d (b lo sso m a n d sh o o t) bud r a t h e r th a n
a sh o o t bud.

A p p ro x im a te ly fo u r o r s ix w e e k s a f te r s c io n g ro w th began,

t r e e s sh o w in g p o o r v ig o r o r c o n ta in in g m ix e d b u d s w e re r e p la c e d by m o re
n o r m a l p la n ts s im i l a r l y g ro w n f o r s u c h a p u r p o s e .
In E x p e rim e n t 2, th e d e s ir e d is o to p ic p h o s p h o ru s w as p la c e d in 0. 2
p e r c e n t o r th o - p h o s p h o r ic a c id c a r r i e r , an d th e pH of the s o lu tio n w as a d ­
ju s te d to 2. 0 w ith p o ta s s iu m h y d ro x id e .

F ifte e n d ro p s of th e is o to p ic t r e a t ­

in g s o lu tio n (30 u c /m l) w e r e a p p lie d to th e f o u rth a n d fifth le a v e s of the

^ M in n e a p o lis H o neyw ell, Brown I n s tr u m e n ts , I n d u s tr ia l D iv isio n ,
P h ila d e lp h ia , P e n n s y lv a n ia .
2
W a rd s N a tu r a l S c ie n c e E s ta b lis h m e n t, 3000 R idgew ood E a s t, R o c h e s te r ,
N ew Y o rk .

34

s c io n .

T h e r e w e r e no v is u a l s y m p to m s w h ich w ould in d ic a te le a f in ju ry .
P la n ts w e r e h a r v e s t e d 24, 48 a n d 96 h o u r s fo llo w in g tr e a tm e n t,

a n d s e p a r a t e d a s fo llo w s: (a) t r e a t e d le a v e s , (b) sh o o t ab o v e t r e a t e d le a v e s ,
(c) sh o o t b elo w t r e a t e d le a v e s , (d) fiv e c e n tim e te r p h lo e m s e g m e n t c o n ­
ta in in g th e b u d union, a n d (e) r o o ts .
In r o o t a b s o rp tio n s tu d ie s of P

( E x p e rim e n t 3) th e t r e a t in g s o lu ­

tio n w a s p r e p a r e d by a d d in g p h o s p h o ru s la b e le d o r th o - p h o s p h o ric a c id to
d i s tille d w a te r so th a t te n m i l l i l i t e r s w ould d e liv e r 108. 5 m i c r o c u r i e s of
P 32 .

T e n m i l l i l i t e r s of th e t r e a t i n g s o lu tio n w e re th e n ad d e d to e a c h c u ltu r e

j a r w ith a r e s u l t i n g s p e c ific a c tiv ity of one m ic r o c u r ie o f P 32 to one m illig r a m
of P
of P

31
32

.

T h e s tio n s w e r e h a r v e s te d 24, 48 a n d 96 h o u rs a f t e r th e a d d itio n
to th e r o o t m e d iu m , a n d s e p a r a te d in to th e fo llo w in g s a m p le s :

(a) sc io n , (b) fiv e c e n tim e te r s e g m e n t c o n ta in in g th e bud union, (c) s to c k
b elo w th e bud union, a n d (d) r o o ts .
R a d io c a lc iu m in w eak h y d r o c h lo ric a c id w a s a p p lie d c a r r i e r f r e e to
th e n u tr ie n t m e d iu m (E x p e rim e n t 4) a t th e r a t e of 0. 5 m ic r o c u r ie of C a^^
to one m illig r a m of C a 40 .

T h e s tio n s w e re h a r v e s te d 48, 96 a n d 144 h o u r s

a f t e r tr e a t m e n t by s e p a r a tin g in to (a) sc io n , (b) bud union, an d (c) s to c k
b elo w th e b u d u n ion.

T h e r e w as no v is ib le in ju ry a p p a r e n t a s a r e s u l t of

t r e a t m e n t of th e r o o t m ed iu m w ith e i th e r P
c o n d itio n s a n d t im e s s tu d ie d .

o r Ca

u n d e r th e d e s c r ib e d

35

W i t h i n t h e s h o r t d u r a t i o n (4 o r

6

w e e k s ) of t h e e x p e r i m e n t s l i t t l e

d i f f e r e n c e s w e r e v i s i b l e in t h e v i g o r of t h e s c i o n e x c e p t on E M XVI.

In

E x p e r i m e n t s 2 a n d 3 M c I n t o s h on E M XVI w a s s l o w e r to i n i t i a t e r o o t s ,
b r e a k b u d a n d p r o d u c e d t h e l e a s t t e r m i n a l g r o w t h ( T a b l e I).

R adioactive

a n a l y s i s of t h e s c i o n s t e m a b o v e t h e t r e a t e d l e a v e s , f o l l o w i n g f o l i a r a p p l i c a t i o n of P
rootstock.

32

, s h o w e d no s i g n i f i c a n t d i f f e r e n c e s i r r e s p e c t i v e of t h e

H o w e v e r , s c i o n s on E M XVI c o n t a i n e d m o r e P

on E M VII, IX o r D e c l i c i o u s s e e d l i n g r o o t s t o c k s .

32

than those

C om parative am ounts

of f o l i a r a p p l i e d r a d i o p h o s p h o r u s r e c o v e r e d in t h e s c i o n s t e m p o r t i o n
i m m e d i a t e l y a b o v e th e b u d u n io n a n d b e l o w t h e t r e a t e d l e a v e s a r e i l l u s ­
t r a t e d in F i g u r e 3 (T o p ) .

T h e q u a n t i t y r e c o v e r e d in t h i s s e g m e n t c l o s e l y

p a r a l l e l e d t h e s c i o n v i g o r o b s e r v e d in th e f i e l d .
P

32

L e s s of th e f o l i a r a p p l i e d

w a s r e c o v e r e d in t h i s s e g m e n t on d w a r f i n g s t o c k s E M VII a n d IX th a n

v i g o r o u s s t o c k s E M XVI a n d D e c l i c i o u s s e e d l i n g .

T h e r e w a s no e x t e n s i v e

i n c r e a s e of th e i s o t o p e in t h i s r e g i o n f r o m 24 to 96 h o u r s a f t e r t r e a t m e n t .
T h i s s u g g e s t e d t h a t t h e b u d u n io n d i d not o f f e r a n o b s t r u c t i o n to the d o w n ­
w a r d t r a n s p o r t of f o l i a r a p p l i e d p h o s p h o r u s .

W it h M c I n t o s h s c i o n on

E M XVI, t h e d e c r e a s e in P 3 2 a t 48 h o u r s , a s c o m p a r e d w it h 24 h o u r s , in
th e s t e m i m m e d i a t e l y a b o v e t h e b u d u n io n w a s c o r r e l a t e d w it h a h i g h e r
c o n c e n t r a t i o n of P 3 2 in t h e s t e m s e g m e n t a b o v e th e t r e a t e d l e a v e s .

This

d e c r e a s e w a s e v i d e n t in a l l s a m p l e s b e l o w t h e t r e a t e d l e a v e s in t h i s r o o t stock.

36

TABLE I
L E N G T H O F T E R M I N A L G R O W T H O F M C I N T O S H SCIONS F O U R AND
SIX W E E K S A F T E R BUD BREAK ON D W A R F I N G AND V IG O R OUS
R O O T S T O C K S R E C O R D E D IN T H R E E E X P E R I M E N T S

M cIntosh Scion
on

Experim ent
31

2^

41

(centim eters)
E M VII

26. 5

15.

E M IX

21. 3

12. 3

E M XVI

15.

D elicious Seedling

23. 1

15. 2

-------

L . S . D. 5%

4. 3

3. 0

3. 4

1%

5. 9

4. 0

4. 1

1

2

8

8

9. 9

M e a s u r e m e n t taken 4 w e ek s a f te r bud b re a k .
M e a su rem en t taken

6

w e e k s a f t e r bu d b r e a k .

13. 1
18. 4
18.

2

Figure 3
R a d i o p h o s p h o r u s (P

32

c p m / g m d r y w t . ) r e c o v e r e d in v a r i o u s
32

t i s s u e s a t 24, 48 a n d 96 h o u r s f o l l o w i n g f o l i a r a p p l i c a t i o n of P oz> l a b e l e d
o r t h o - p h o s p h o r i c a c i d ( 0 . 2 % ) to M c I n t o s h s c i o n s b u d d e d on E a s t M a i l i n g
(EM) VII, IX a n d XVI a n d D e l i c i o u s s e e d l i n g r o o t s t o c k s .
(Top) T h e M c I n t o s h s c i o n s e g m e n t i m m e d i a t e l y a b o v e the
bud union.
(Bottom) T h e five c e n t i m e t e r p h l o e m s e g m e n t c o n t a i n i n g
the bud union.

I IO O

1000

-------------------

300

CPM/GM

DRY

WT

500

SEEDLING
EM VII
EM
IX
EM XVI
1 0 0

1000

500

300
1 0 0

24

48
HOURS

FOLLOWING

96
T R E A T ME NT

38

T h e q u a n t i t y of f o l i a r a b s o r b e d P

on

r e c o v e r e d in th e p h l o e m s e g ­

m e n t c o n t a i n i n g t h e b u d u n io n f o r m e d by M c I n t o s h s c i o n s on E M VII, IX
a n d XVI a n d D e l i c i o u s s e e d l i n g r o o t s t o c k s d i d no t d i f f e r s i g n i f i c a n t l y a m o n g
r o o t s t o c k s ( T a b l e II).

At 48 h o u r s a ll , w i t h t h e e x c e p t i o n of E M XVI, h a d

atta in ed a p la te au and r e m a in e d re la tiv e ly co n stan t u nder th e se conditions
( F i g u r e 3, B o t t o m ) .

It i s i n t e r e s t i n g to n o t e t h a t o nl y in M c l n t o s h / E M IX

does the p h lo e m s e g m e n t co n tain in g the bud union contain m o r e P

32

per

g r a m d r y w e i g h t t h a n t h e s t e m ( F i g u r e 3, T o p ) i m m e d i a t e l y a b o v e it.
T h i s m a y i n d i c a t e t h a t t h e r e i s a n a c c u m u l a t i o n a t t h e b u d union, but if
so, t h e m a g n i t u d e i s v e r y s m a l l w i t h i n t h e f i r s t 96 h o u r s , a n d d o e s not
d i f f e r s i g n i f i c a n t l y f r o m t h e s t e m a b o v e it.
S i g n i f i c a n t l y d i f f e r e n t q u a n t i t i e s of f o l i a r a b s o r b e d P 32 w e r e r e ­
c o v e r e d in t h e r o o t s of t h e v a r i o u s r o o t s t o c k s ( T a b l e III).
a n a l y s i s r e v e a l e d t h a t t h e l e a s t a m o u n t of P

32

Radioactive

w a s t r a n s p o r t e d to th e

r o o t s of E M IX a n d XVI, t h e g r e a t e s t q u a n t i t y in D e l i c i o u s s e e d l i n g s
w i t h i n t e r m e d i a t e v a l u e s f o r E M VII.

The fo liar applied P

32

transported

to t h e r o o t s of E M IX i n c r e a s e d p r o p o r t i o n a t e l y w ith t i m e ( F i g u r e 4,
B ott o m ).

E a s t M a i l i n g VII a n d D e l i c i o u s s e e d l i n g a c c u m u l a t e d P

g r e a t e r r a t e d u r i n g t h e f i r s t 48 h o u r s a n d t h e n l e v e l e d off.
XVI a g a i n s h o w e d l e s s a c t i v i t y at 48 h o u r s t h a n a t 24 h o u r s .

32

at a

While EM
No e x p l a n a ­

t i o n c a n be o f f e r e d to a c c o u n t f o r t h e b e h a v i o r of E M XVI a t 48 h o u r s .

39

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F ig u re 4
R a d i o p h o s p h o r u s (P

QO

c p m / g m d r y wt. ) r e c o v e r e d in v a r i o u s

t i s s u e s of M c I n t o s h s c i o n s b u d d e d on E a s t M a i l i n g (EM) VII, IX a n d
XVI a n d D e l i c i o u s s e e d l i n g r o o t s t o c k s , 24, 48 a n d 96 h o u r s fo l l o w i n g
t r e a t m e n t w i t h P 32
(Top) R a d i o p h o s p h o r u s c o n t e n t of M c I n t o s h s c i o n s f o l l o w i n g
a p p l i c a t i o n to th e r o o t m e d i u m .
(Bottom) R a d i o p h o s p h o r u s c o n t e n t of r o o t s f o l l o w i n g f o l i a r a p ­
plication.

700
500
EM

300

XVI

SEEDLING

200
WT

/

E
M

CPM/6M

DRY

_________________

IO O

80
1 0

«•••

,

.•••“

E M IX

«•••

‘.

400

EM VII
E M IX

200

EM

XVI

1 0 0

24

48
HOURS

FOLLOWING

TREATMENT

42

It a p p e a r s f r o m t h e s e d a t a t h a t t h e r o o t s t o c k in s o m e w a y m o d i f i e s th e p h y s i o l o g y of t h e s c i o n .

T h i s m o d i f i c a t i o n p r o b a b l y e x p r e s s e s i t s e l f in th e

f o l i a r a b s o r p t i o n o r t r a n s p o r t m e c h a n i s m to m o r e c l o s e l y a c c o m m o d a t e
th e r e q u i r e m e n t s of the sto c k .

If i t w e r e a q u e s t i o n of r e d u c e d t r a n s p o r t

t h r o u g h t h e s t o c k , t h e n o ne w o u l d e x p e c t a n a c c u m u l a t i o n of t h e f o l i a r a p plied P

32

a b o v e t h e b u d uni o n.

T h i s i s not t h e c a s e , h o w e v e r ,

s i n c e on

a c o m p a r a t i v e b a s i s a s i m i l a r d i s t r i b u t i o n of P 32 e x i s t e d b e t w e e n t h e s t e m
s e g m e n t a b o v e t h e b u d u n io n a n d r o o t s in th e v a r i o u s s t o c k s .
W h e n P 32 w a s a p p l i e d t h r o u g h t h e r o o t s no s i g n i f i c a n t d i f f e r e n c e s
c o u l d be d e t e c t e d in a f i v e - c e n t i m e t e r s t e m s e g m e n t c o n t a i n i n g th e b ud
u n i o n , o n t h e v a r i o u s r o o t s t o c k s ( T a b l e II).
H ow ever, a c l e a r d istinction w as a p p a re n t between the dw arfing
( E M VII a n d IX), a n d v i g o r o u s ( E M XVI a n d D e l i c i o u s s e e d l i n g ) r o o t s t o c k s
in t h e i r a b i l i t y to a b s o r b a n d t r a n s l o c a t e P

32

to th e s c i o n .

The least amount

of P ^ 2 w a s r e c o v e r e d in th e M c I n t o s h s c i o n on E M IX, w i t h a n i n t e r m e d i a t e
v a l u e f o r E M VII a n d t h e g r e a t e s t on D e l i c i o u s s e e d l i n g a n d E M XVI ( T a b l e
III).

At 24 h o u r s t h e m o s t v i g o r o u s c o m b i n a t i o n ( M c l n t o s h / E M XVI) d i f ­

f e r e d f r o m t h e m o s t d w a r f i n g ( M c l n t o s h / E M IX) by only 100 c o u n t s p e r
m inute.

T h e s e d i f f e r e n c e s w e r e a c c e n t u a t e d a t 48 a n d 96 h o u r s a f t e r th e

ad d itio n of the iso to p e to the ro o t m e d iu m .

It i s i n t e r e s t i n g to n o t e t h a t

t h e t i m e a c t i v i t y c u r v e s f o r d w a r f i n g s t o c k s (E M VII a n d IX) w e r e a l m o s t

43

p a r a l l e l ( F i g u r e 4, T o p ).

T h e v i g o r o u s s t o c k s (E M XVI a n d D e l i c i o u s s e e d ­

lin g ) a l s o c l o s e l y p a r a l l e l e d e a c h o t h e r .

The o rd e r of P ^ a b so rp tio n and

t r a n s p o r t in s t i o n s ( M c l n t o s h / E M IX, VII a n d XVI a n d D e l i c i o u s s e e d lin g )
c l o s e l y f o ll o w e d t h e v e g e t a t i v e g r o w th o b s e r v e d in M c I n to s h s c i o n s on E M
IX, VII a n d XV I a n d D e l i c i o u s s e e d l i n g u n d e r c o m m e r c i a l p r a c t i c e .

T hese

d a t a s h o w th a t e a c h s t i o n h a s a c h a r a c t e r i s t i c t i m e a b s o r p t i o n c u r v e , w h ic h
i s c l o s e l y c o r r e l a t e d w ith v i g o r .
W hen Ca

45

w a s a p p l i e d to t h e r o o t m e d iu m o f M c I n to s h s c i o n s on

E M VII, IX a n d XVI r o o t s t o c k s ,

s i g n i f i c a n t l y g r e a t e r q u a n t i t i e s of C a

w e r e r e c o v e r e d in th e M c I n t o s h s c i o n s on E M XVI ( F i g u r e 5).

45

The le ast

q u a n tity of th e r o o t a b s o r b e d C a 45 w a s fou nd in th e s c i o n s on th e d w a r f in g
s t o c k E M IX.
in C a

45

M c I n to s h s c i o n s on E M VII ( s e m i - d w a r f ) w e r e i n t e r m e d i a t e

c o n te n t ( F i g u r e 5).

A s w ith r o o t a b s o r b e d P

52

, th e d w a r f in g r o o t ­

s t o c k s (E M VII a n d IX) e x h i b i t e d s i m i l a r t i m e a b s o r p t i o n c u r v e s .

R ad io ­

a c t i v i t y in th e s c i o n s on th e v i g o r o u s s t o c k (E M XVI) i n c r e a s e d w ith t i m e
a n d l a r g e r a m o u n t s w e r e p r e s e n t t h a n in th e s c i o n s on d w a r f in g s t o c k s
(E M V II a n d IX).

C o n s e q u e n tly , th e i n h e r e n t c a p a c i t y of E M XVI r o o t s t o c k s

to a b s o r b c a l c i u m w a s g r e a t e r t h a n e i t h e r E M VII o r E M IX.
T h e r e w a s no a p p a r e n t r e l a t i o n b e tw e e n s t i o n v i g o r a n d C a
c e n t r a t i o n in th e s t e m s e g m e n t c o n t a i n i n g th e b u d u n io n ( T a b le II).

45

con­

No

m a r k e d a c c u m u l a t i o n o f r o o t a p p l i e d C a ^ w a s o b s e r v e d a t th e bu d u n io n
( T a b l e II), i n d i c a t i n g t h a t th e u n io n d id n o t m a t e r i a l l y i m p e d e C a
d u r i n g th e t i m e s s t u d i e d .

45

tra n sp o rt

F ig u re 5
R a d i o c a l c i u m ( C a ^ c p m / g m d r y w t . ) d e t e c t e d in th e
M c I n to s h s c i o n s on E M VII, IX a n d XVI r o o t s t o c k s , 48, 96 a n d
144 h o u r s f o llo w in g i t s a d d itio n to th e r o o t m e d i u m .

5000

EM

XVI

EM

VII

EM

IX

500

CPM/GM

DRY

WT

1000

100

50

30
48
HOURS

96
FOLLOWING

144
TREATMENT

45

E x p e rim e n t 5

T h e t r a n s p o r t of f o l i a r a p p lie d r a d io p h o s p h o ru s by g e n e tic a lly
d w a rfe d a n d n o n - d w a rf e d to m a to p la n ts w a s s tu d ie d .

S e e d lin g s of g e n e ­

tic a lly d w a rf ( v a r. M a c A r th u r 741) a n d n o n -d w a rf (v a r. S to k e sd a le ) to m a to
p la n ts w e r e s t a r t e d in v e r m ic u lite a n d th e n t r a n s f e r r e d a s s m a ll s e e d lin g s
to s o lu tio n c u l t u r e s u s in g a s ta n d a r d H o ag lan d fo rm u la tio n .
b lo c k d e s ig n w ith fiv e r e p lic a tio n s w as u se d .

A r a n d o m iz e d

A f te r te n d ay s, o n e - h a lf of

th e p la n ts of e a c h v a r i e t y w e r e t r e a t e d by a p p ly in g 0. 1 m ic r o c u r ie of P
to th e f i r s t t r u e le a f.

32

A t th e tim e of tr e a tin g , both d w a rf an d v ig o ro u s

p la n ts h a d th e s a m e n u m b e r (4 -5 ) of le a v e s , th u s t h e i r p h y s io lo g ic a l d e ­
v e lo p m e n t w a s c o m p a r a b le .
P la n ts w e r e h a r v e s te d a f t e r 6 a n d 24 h o u rs an d s e p a r a te d into th e
fo llo w in g p a r t s : (a) t r e a t e d le a f, (b) s te m p lu s n o n - tr e a te d le a v e s , a n d
(c) r o o ts .

T h e s a m p le s w e re p la c e d in 50 m i l l i li t e r b e a k e r s , d r ie d in a

f o r c e d a i r o v en (70*C) f o r 96 h o u r s , a n d th e n c o u n te d d ire c tly .

No c o r ­

r e c t i o n w a s m a d e f o r s e lf a b s o rp tio n .
09

T h e to ta l P

a b s o r b e d a n d tr a n s lo c a te d fro m th e t r e a t e d le a f w as

a lm o s t id e n tic a l f o r th e d w a rf (1627. 7 cpm ) a n d n o n -d w a rf (1626. 5 cpm )
to m a to p la n ts (T a b le IV).

T h is in d ic a te d th a t no d iff e r e n c e s e x is te d b etw een

th e d w a rf a n d n o n - d w a rf to m a to p la n ts in t h e ir a b ility to a b s o r b f o lia r

46

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47

a p p lie d r a d io p h o s p h o r u s .

H o w e v e r, th e r e w e r e s ig n ific a n t d iff e r e n c e s in th e

d is tr ib u tio n o f th e a b s o r b e d p h o s p h o ru s .

M o re P

oo

w as r e c o v e r e d in th e

s te m s a n d n o n - tr e a t e d le a v e s o f th e d w a rf th a n th e n o n -d w a rf p la n ts , but
th e d if f e r e n c e s w e r e n o t s ig n if ic a n t (T a b le IV).
f o lia r a p p lie d P

32

S ig n ific a n tly m o re of th e

w a s r e c o v e r e d in th e r o o ts of th e n o n -d w a rf th a n th e d w a rf

to m a to p la n ts , in d ic a tin g th a t l e s s f o lia r a p p lie d P*^ w as tr a n s lo c a te d to th e
r o o t o f th e d w a rf to m a to p la n t.

T h u s , w ith to m a to e s , the a b s o rp tio n c a p a ­

c ity o f le a v e s of d w a rf a n d n o n -d w a rf p la n ts d id not d iffe r, but t r a n s p o r t to
th e r o o ts o f th e d w a rf p la n t w as r e d u c e d .

E x p e rim e n t 6
P r e v io u s s tu d ie s (E x p e r im e n t 5) in d ic a te d th a t d w a rf an d n o n -d w a rf
to m a to p la n ts a b s o r b e d e q u a l q u a n titie s of f o lia r a p p lie d P

32

, but le s s P

t r a n s p o r t e d to th e r o o ts o f th e d w a rf th a n th e n o n -d w a rf p la n t.

32

w as

T he follow ing

e x p e r im e n t w a s d e s ig n e d to o b s e rv e th e a b s o rp tio n an d su b se q u e n t tr a n s p o r t
of f o li a r a p p lie d P

QO

in c h e m ic a lly d w a rfe d an d n o n -d w a rfe d to m a to p la n ts .

T o m a to ( v a r. S to k e sd a le ) s e e d lin g s w e r e g ro w n a s in th e p r e v io u s
e x p e r im e n t.

A f te r te n d a y s of g ro w th in s ta n d a r d H oagland so lu tio n c u ltu r e s ,

th e fo lia g e w a s d ip p e d in s o lu tio n s (0, 1000 o r 4000 ppm ) of m a le ic h y d ra z id e
( fo r m u la te d a s a w a te r s o lu b le d ie th a n o la m in e s a lt of 1, 2 - d ih y d r o - 3, 6 - p y r id a z in e d io n e ).

F o u r d a y s a f t e r t r e a tm e n t w ith m a le ic h y d ra z id e , 0. 1 m ic r o c u r ie

48

of P

32

v e ste d

( a s H ^P
6

32

O 4 ) w a s a p p l i e d to th e f i r s t t r u e l e a f .

The p la n ts w ere h a r ­

a n d 24 h o u r s l a t e r , a n d s a m p l e s p r e p a r e d a n d c o u n t e d a s d e s c r i b e d

in th e p r e v i o u s e x p e r i m e n t .
N e w to p g r o w t h w a s g r e a t l y r e d u c e d in t h e m a l e i c h y d r a z i d e t r e a t e d
p la n ts.

T o m a t o p l a n t s t r e a t e d w ith 4 0 0 0 p a r t s p e r m i l l i o n s h o w e d b u r n i n g

o f th e l e a f e d g e s .

H o w e v e r , no i n j u r y o c c u r r e d w ith 1000 p a r t s p e r m i l l i o n .

T h e p e r c e n t o f th e f o l i a r a p p l i e d P

09

r e c o v e r e d in th e s t e m o r r o o t s d id n o t

v a r y s i g n i f i c a n t l y w ith c h e m i c a l t r e a t m e n t a t s i x h o u r s .

S ig n if ic a n tly l e s s

f o l i a r a p p l i e d r a d i o p h o s p h o r u s w a s f o u n d in th e s t e m s a n d n o n - t r e a t e d l e a v e s
a t 24 h o u r s o f p l a n t s t r e a t e d w ith 4 0 0 0 p a r t s p e r m i l l i o n of m a l e i c h y d r a z i d e .
No d i f f e r e n c e s w e r e o b s e r v e d in th e P
t r e a t m e n t ( T a b l e V).

on

c o n te n t of th e r o o t s due to c h e m i c a l

T h e t o t a l a m o u n t of f o l i a r a p p l i e d P 3^ a b s o r b e d a n d

t r a n s p o r t e d o u t o f th e t r e a t e d l e a f d id n ot d i f f e r s i g n i f i c a n t l y w ith c h e m i c a l
t r e a t m e n t a t s i x h o u r s ( T a b l e V), h o w e v e r , a t 24 h o u r s c o n s i d e r a b l y l e s s
P 3 2 w a s t r a n s p o r t e d o u t o f t h e t r e a t e d l e a f in to m a to p l a n t s t r e a t e d w ith
400 0 p a r t s p e r m i l l i o n o f m a l e i c h y d r a z i d e .

T h i s i n d i c a t e d th a t c h e m i c a l l y

d w a rfe d to m a to p la n ts a b s o r b e d le s s fo lia r a p p lie d P
t h e a b s o r b e d p ^ 2 f r o m t h e t r e a t e d le a f, o r b o th .

32

, t r a n s p o r t e d l e s s of

49

TABLE V
T R A N S P O R T O F F O L I A R A P P L IE D R A DIO PH O SPH O R U S (P32) IN C H E M IC A L L Y
D W A R F E D 1 T O M A T O P L A N T S (VAR. S T O K E S D A L E )

M ale ic H y d ra z id e
C o n ce n tra tio n

T im e A fte r T r e a tm e n t
—--------------------------------------------------------------------------------------6 H o u r s __________________
24 H o u r s _______
S te m
Root
T o tal
S te m
Root
T o tal

(p p m )

(m e a n s of

re p lic atio n s - % reco v ered )

. 42

. 24

.

1 0 0 0

.4 0

.

20

. 60

4 00 0

. 39

. 30

. 69

0

(c o n tro l)

8

66

. 79

.4 9

2

1

. 56

. 39

1. 95

. 42

. 95

. 53

2

1M a l e i c h y d r a z i d e a p p l i e d 4 d a y s p r i o r to t r e a t m e n t w ith P ° A
O
D iffe rs s ig n ific a n tly at

1

% lev el fro m co n tro l.

. 28

1

50

S e c tio n B
E f f e c t o f t h e G r a f t U n io n a n d S c io n O r i e n t a t i o n
on N u t r i e n t T r a n s p o r t

E x p e rim en t 7
T h e e f f e c t o f t h e g r a f t u n io n on th e t r a n s p o r t of p h o s p h o r u s w a s s t u d i e d by
fo llo w in g t h e a c r o p e t a l m o v e m e n t of P

09

un d er sh o rt tim e in te rv a ls .

V ig o r o u s

t o m a t o p l a n t s ( v a r . S t o k e s d a l e ) w ith 8 - 1 0 l e a v e s , g r o w n in s o i l c u l t u r e s ,
w e re sa d d le g ra fte d .

T h e s c i o n c o n s i s t e d of a t w o - i n c h s t e m p i e c e w ith one

g r o w i n g p o in t in t h e l e a f a x i l .

T o m i n i m i z e p l a n t v a r i a b i l i t y th e s c i o n w a s

o f th e s a m e c l o n e a s th e s to c k .

F o r c o n tro ls, o th e r p la n ts w e re d e ca p ita te d

a t a p p r o x i m a t e l y th e s a m e h e ig h t a s th e top o f th e s c i o n of th e g r a f t e d p l a n t s .
I m m e d i a t e l y a f t e r g r a f t i n g , th e p l a n t s w e r e p l a c e d in a h u m id ity (9 5 -1 0 0 % R. H . )
c h a m b e r for fo u r days.

T h e y w e r e t h e n g r o w n in a g r e e n h o u s e u n d e r th e

n o r m a l l y lo n g d a y s of l a t e s p r i n g a n d e a r l y s u m m e r w ith h igh n u t r i e n t l e v e l s
in th e r o o t m e d i u m .

F i f t y - t w o d a y s a f t e r g r a f t i n g , 30 m i l l i l i t e r s o f a . 05 M

p o t a s s i u m d i - h y d r o g e n p h o s p h a t e s o l u t i o n c o n t a i n i n g one m i c r o c u r i e of P

32

p e r m i l l i l i t e r w a s a p p l i e d to t h e s o i l in a s m a l l t r e n c h a r o u n d th e b a s e of
e a c h p la n t.
T h e p l a n t s w e r e h a r v e s t e d 3, 12, 24, 48 a n d 96 h o u r s fo llo w in g th e a d ­
d itio n o f P 3 2 to th e r o o t m e d i u m .

T h e g r a f t e d p l a n t s w e r e s e p a r a t e d into th e

s c i o n a n d g r a f t u n io n .

In th e n o n - g r a f t e d p l a n t s , th e n e w s h o o t w a s h a r ­

v e s t e d a s a s c i o n a n d a c o m p a r a b l e s t e m s e g m e n t a s th e g r a f t un ion .
D r i e d p l a n t t i s s u e s a m p l e s w e r e g r o u n d to p a s s th r o u g h a 40 m e s h
s c r e e n a n d t h e n w e t a s h e d p r i o r to r a d i o a c t i v e a s s a y .

E ach sam p le w as

c o u n t e d t h r e e t i m e s a n d t h e a v e r a g e c o u n t p e r m i n u t e p e r g r a m d r y w e ig h t
of tis s u e r e c o r d e d .
R a d i o p h o s p h o r u s (P

32

) r e c o v e r e d in th e s c i o n s a n d g r a f t u n io n s of

g r a f t e d p l a n t s a n d c o r r e s p o n d i n g t i s s u e s (new s h o o t a n d s t e m s e g m e n t
c o m p a r a b l e to th e g r a f t u n io n s e g m e n t ) of n o n - g r a f t e d p l a n t s a r e sh ow n in
T a b l e VI.

T h e r a d i o a c t i v i t y fo u n d in th e s c i o n s a n d g r a f t u n io n s g e n e r a l l y

i n c r e a s e d w ith t i m e in b o th g r a f t e d a n d n o n - g r a f t e d p l a n t s .

No e x p la n a tio n

is o f f e r e d f o r th e lo w r a d i o a c t i v i t y of n o n - g r a f t e d s c i o n s at 24 h o u r s a n c th e
g r a f t e d s c i o n s a t 48 h o u r s .

T h e r e w e r e no c o n s i s t e n t d i f f e r e n c e s in th e

P 32 c o n c e n t r a t i o n o f th e s c i o n s o r g r a f t u n io n s of g r a f t e d o r c o m p a r a b l e
p o r t i o n s o f n o n - g r a f t e d p l a n t s a t a n y of th e t i m e i n t e r v a l s a f t e r th e is o to p e
w a s a d d e d to th e r o o t m e d i u m .

L i k e w i s e , no s ig n if ic a n t d i f f e r e n c e s w e r e

o b s e r v e d b e tw e e n th e m e a n s f o r a l l t i m e s (3, 12, 24, 48 a n d 96 h o u r s ) f o r
g ra fte d v e r s u s n o n -g r a fte d to m a to p la n ts .

No o b s t r u c t i o n , a s s o c i a t e d w ith

g r a f t i n g , to th e u p w a r d t r a n s p o r t of r o o t a b s o r b e d r a d i o p h o s p h o r u s w a s o b ­
served.

F u r t h e r m o r e , th e p r e s e n c e o r a b s e n c e of th e g r a f t u n io n in t o m a to

p l a n t s d id not a f f e c t th e q u a n t i t y of r a d i o p h o s p h o r u s a b s o r b e d a n d t r a n s p o r t e d
to th e s c io n .

52

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53

E x p e rim en t

8

N u t r i e n t d i s t r i b u t i o n in g r a f t e d t o m a t o p l a n t s w a s a s c e r t a i n e d by
th e a u to ra d io g r a p h ic te c h n iq u e .

S ix w e e k o ld t o m a t o p l a n t s ( v a r . M o r e t o n

H y b rid ) w e r e g r a f t e d a s d e s c r i b e d in th e p r e v i o u s e x p e r i m e n t .

A fte r fo u r

w e e k s , a s o l u t i o n o f th e d e s i r e d i s o t o p e w a s a p p l i e d to th e s o il in a b a n d
a r o u n d th e b a s e of e a c h p l a n t .
Fe
P

55-59

32

Mo

, Zn

65

, Rb

86

, Sr

89

T he iso to p e s; N a22, C l36, C a 45, M n5 2 -5 4 ,

a n d Ba

a s o r t h o - p h o s p h o r i c a c id , S

99

a s a m m o n iu m m o ly b d a te .

re c eiv e d

1 0 0

35

140

w e r e u s e d in t h e i r c h l o r i d e f o r m ,

a s su lfa te, Cu

64

as co p p er n itra te , and

T h e s o i l in w h ic h e a c h p l a n t w a s g r o w in g

m i c r o c u r i e s o f o n e o f th e r e s p e c t i v e i s o t o p e s e x c e p t in the

c a s e o f c h l o r i n e w h e r e 15 m i c r o c u r i e s w e r e u s e d .
T h e p l a n t s w e r e h a r v e s t e d 48 h o u r s f o llo w in g th e a d d itio n of th e
r a d i o a c t i v e i s o t o p e s to th e r o o t m e d i u m .

In h a r v e s t i n g , a 2 0 - c e n t i m e t e r

s e g m e n t of th e s t e m m i n u s a p p e n d a g e s ( e x c e p t in th e c a s e o f M o

99

th e a p ­

p e n d a g e s w e r e r e t a i n e d ) b u t i n c lu d in g th e g r a f t union, w a s r e m o v e d a n d
p r e s s e d b e t w e e n b o t a n i c a l d r y i n g p a p e r in a f o r c e d a i r o v e n a t 70° C.
d r y in g , th e s a m p l e s w e r e t r a n s f e r r e d to new

8

x

10

U pon

in c h s h e e t s of b o ta n ic a l

d r y i n g p a p e r , c o v e r e d w ith S a r a n w r a p (Dow C h e m i c a l C o m p a n y , M id la n d ,
M i c h i g a n ) a n d e x p o s e d to K o d a k X - r a y film (Blue b r a n d ) .

T h re e stem s p e c i­

m e n s fro m t h r e e p la n ts re s p e c tiv e ly , fo r e a c h ra d io iso to p e , w e re u tiliz e d
f o r p r o c u r i n g th e a u t o r a d i o g r a m s .

54

In o r d e r to o b t a i n a d e s i r a b l e i m a g e th e p r e s s e d a n d d r i e d s t e m
s e c t i o n s w e r e e x p o s e d a s fo llo w s :
(a) 24 H o u r s

N a 2 2 , P 3 2 , C l 3 6 , Ca.4 5 , R b 8 6 , S r 8 9 , M o 9 9

(b)

S3 5

7 D ays

(c) 15 D a y s

M n 5 2 ' 5 4 ,C u 6 4

(d) 21 D a y s

F e * ^ ~ " ,Z n ^

Ba1 4 0

A ll a u t o r a d i o g r a m s s h o w a s i m i l a r i t y in t h a t t h e r e w a s no a c c u m u l a t i o n
of th e i s o t o p e a t th e s i t e o f th e g r a f t u n io n ( F i g u r e s
la t i o n o f a n y of t h e i s o t o p e s a b o v e o r b e lo w
p o s s ib le e x c e p tio n of c h lo rin e ( C l ^ ) .

6

, 7 , 8 , 9).

th e g r a f t u n io n

No a c c u m u ­

o c c u r r e d w ith th e

T h i s w a s of l i t t l e s i g n i f i c a n c e

s in c e

c h l o r i n e t e n d s to a c c u m u l a t e in th e v a s c u l a r s y s t e m of to m a to p l a n t s
( B r o y e r 1956).

E x p e rim en t 9

D is trib u tio n of ro o t a b s o r b e d ra d io c a lc iu m w as next stu d ie d th ro u g h
a d w a r f i n g a p p le i n t e r s t e m in v o lv in g tw o g r a f t u n io n s .

T h re e tw o -y e a r-o ld

M c I n t o s h t r e e s * c o n t a i n i n g a C l a r k ’s d w a r f i n t e r s t e m w e r e s t a r t e d in s a n d
c u l t u r e s w ith s t a n d a r d H o a g la n d s o lu tio n .

T w o w e e k s a f t e r th e i n i t i a t i o n

o f b u d a c t i v i t y , a n d w h ile th e t r e e s w e r e a c t i v e l y g r o w in g ,
o f C a 4 "* w e r e a p p l i e d to th e r o o t m e d i u m o f e a c h t r e e .

100

m ic ro c u rie s

A 20 c e n t i m e t e r

T r e e s s u p p l i e d by S t a r k B r o t h e r s N u r s e r y , L o u i s i a n a , M i s s o u r i .

Figure 6
A u t o r a d i o g r a m s of g r a f t e d t o m a t o s t e m s s h o w in g th e d i s t r i ­
b u tio n of C a 4 ^ (left), Ba 4 4 *4 ( c e n t e r ) , a n d S r " ( r ig h t) , 48 h o u r s fo llo w ­
in g th e a d d itio n of th e i s o t o p e s to th e r o o t m e d i u m .
s i t e of th e g r a f t u nio n).

( A r r o w s i n d ic a te

55

Figure 7
A u t o r a d i o g r a m s of g r a f t e d t o m a t o s t e m s s h o w in g th e d i s t r i ­
b u tio n of C l 3^ (left), S35 ( c e n t e r ) a n d N a ^ ( r ig h t) , 48 h o u r s f o llo w in g
th e a d d itio n of th e i s o t o p e s to th e r o o t m e d i u m .
of th e g r a f t u n ion ).

( A r r o w s i n d i c a t e s ite

56

Figure 8
A u t o r a d i o g r a m s of g r a f t e d t o m a t o s t e m s i l l u s t r a t i n g th e d i s t r i ­
b u tio n of

(le ft),

s o il a p p l i c a t i o n .

( c e n t e r ) a n d M n ^ ' ^ ( r ig h t) , 48 h o u r s fo llo w in g

( A r r o w s i n d i c a t e s i t e of th e g r a f t u n io n ).

57

F igure 9
A u t o r a d i o g r a m s of g r a f t e d t o m a t o s t e m s i l l u s t r a t i n g th e d i s t r i b u tio n of Z n
m e d iu m .

65

(le ft) a n d M o

99

( r ig h t) , 48 h o u r s a f t e r t r e a t i n g th e r o o t

( A r r o w s i n d i c a t e s i t e of th e g r a f t u n io n ).

58

59

s e g m e n t o f th e s t e m c o n t a i n i n g th e i n t e r s t e m w a s r e m o v e d 24 h o u r s a f t e r
tre a tin g .

T he s e g m e n ts w e re d rie d , se c tio n e d and a u to ra d io g ra m s p r e p a r e d .
T h e d w a r f i n g i n t e r s t e m w a s l a r g e r in d i a m e t e r th a n e i t h e r th e

s t o c k o r irie s c i o n ( F i g u r e 10).

T h e g r a f t u n io n s b e tw e e n th e s c i o n a n d i n t e r ­

s t e m w e r e w e ll f o r m e d , w h e r e a s t h o s e b e tw e e n th e i n t e r s t e m a n d s t o c k
s h o w e d e v i d e n c e of i n t e r n a l b r e a k d o w n .
e v id e n c e of s t r u c t u r a l w e a k n e s s .

T h e r e w a s , h o w e v e r , no e x t e r n a l

T h e p r i m a r y a n d i n i t i a l p a th w a y f o r th e

t r a n s p o r t of r o o t a b s o r b e d r a d i o c a l c i u m d u r i n g th e f i r s t 24 h o u r s a p p e a r e d
to be th e n e w ly f o r m e d x y le m ( A u t o r a d i o g r a m , F i g u r e 10).

No i n t e r r u p t i o n

of t r a n s p o r t w a s n o te d , b u t f o r c o n c l u s i v e e v id e n c e s e r i a l s e c t i o n s w o u ld
be n e c e s s a r y .

E x p e r i m e n t 10

D u r i n g th e c o u r s e of e a r l i e r e x p e r i m e n t a t i o n it w a s o b s e r v e d t h a t
Ca

45

f a i l e d to m o v e out of th e t r e a t e d le a f .

F u r t h e r m o r e , w hen a p p l i e d to

. i-

th e s t e m C a

45

m o v e d o n ly a c r o p e t a l l y .

T h e ’’p o l a r " t r a n s p o r t of C a

, in

09

c o m p a r i s o n w ith P

, w a s n e x t s t u d i e d by u t i l i z i n g g r a f t u n io n s o f to m a to

p l a n t s in v o lv in g r e v e r s e d p o l a r i t y .
U n if o r m f i v e - w e e k - o l d t o m a t o p l a n t s ( v a r . S to k e s d a l e ) w e r e d e c a p i ­
t a t e d a b o v e th e s e c o n d t r u e le a f .

T w o l a t e r a l s w e r e p e r m i t t e d to g r o w a n d

f o u r w e e k s l a t e r th e l o w e r l a t e r a l o f o n e p l a n t w a s g r a f t e d (a p e x to a p ex)
in to th e u p p e r l a t e r a l of a n o t h e r p l a n t ( F i g u r e 11).

F i g u r e 10
P h o to g r a p h (le ft) i l l u s t r a t i n g a l o n g itu d in a l s e c t i o n t h r o u g h a
C l a r k ’s d w a r f i n t e r s t e m w ith a M c I n to s h s c i o n a n d H i b e r n a l s to c k , a n d
a u t o r a d i o g r a m ( r ig h t) s h o w in g th e d i s t r i b u t i o n a n d p a th w a y of t r a n s p o r t
of Ca

45

, 24 h o u r s a f t e r a p p l i c a t i o n to th e r o o t m e d i u m .

F i g u r e 11
A u n io n f o r m e d by g r a f t i n g t o g e t h e r (a p e x to a p e x ) th e
f o r c e d l a t e r a l s of tw o t o m a t o p l a n t s .

61

62

T h r e e w e e k s a f t e r g r a f t i n g , a n d w h e n th e f i r s t f l o w e r s w e r e op en ,
100 m i c r o c u r i e s o f C a

45

o r 50 m i c r o c u r i e s o f P

32

w e r e a d d e d to th e s o il in

w h ic h th e r o o t s o f o n e of th e g r a f t e d p l a n t s w e r e g r o w in g .

The g ra fte d

l a t e r a l s w e r e s e v e r e d f r o m th e tw o p l a n t s 6 o r 12 d a y s a f t e r t r e a t m e n t
a n d s e g m e n t e d a s fo llo w s : (a) f iv e c e n t i m e t e r s e g m e n t c o n t a i n i n g th e g r a f t
u n io n , (b) fiv e c e n t i m e t e r s e g m e n t a d j a c e n t to t h e g r a f t u n io n a n d f r o m th e
p l a n t r e c e i v i n g th e r a d i o a c t i v e is o to p e , a n d (c) fiv e c e n t i m e t e r s e g m e n t a d ­
j a c e n t to th e u n io n b u t f r o m th e p l a n t r e c e i v i n g no r a d i o a c t i v e is o to p e .
T w e lv e d a y s f o ll o w i n g t r e a t m e n t y o u n g i m m a t u r e f r u i t s w e r e h a r v e s t e d f r o m
b o th p l a n t s .

S te m s e g m e n t a n d f r u i t s a m p l e s w e r e p r e p a r e d a n d a s s a y e d

fo r ra d io a ctiv ity .

T h r e e a p e x to a p e x u n io n s w ith a p p r o a c h i n g l a t e r a l s w e r e

u s e d to o b t a i n a r e p r e s e n t a t i v e a u t o r a d i o g r a m ( F i g u r e 12).
V a l u e s f o r r a d i o a c t i v i t y in th e g r a f t e d l a t e r a l s a n d y o u n g f r u i t s
a r e g iv e n in T a b l e V II.

For P

32

no s i g n i f i c a n t d i f f e r e n c e s w e r e o b s e r v e d

b e t w e e n th e fiv e c e n t i m e t e r l a t e r a l s e g m e n t on th e t r e a t e d s id e a n d th e g r a f t
u n io n s e g m e n t , o r b e t w e e n t h e g r a f t u n io n s e g m e n t a n d th e fiv e c e n t i m e t e r
l a t e r a l s e g m e n t on th e n o n - t r e a t e d s id e .

H o w e v e r , a s ig n i f i c a n t l y h i g h e r

v a lu e w a s o b s e r v e d f o r th e fiv e c e n t i m e t e r l a t e r a l s e g m e n t on th e t r e a t e d
s id e t h a n f o r th e c o m p a r a b l e s e g m e n t on t h e n o n - t r e a t e d s i d e .

R ad io p h o s­

p h o r u s w a s a l s o r e c o v e r e d in t h e i m m a t u r e f r u i t s o c c u r r i n g on th e s e c o n d
l a t e r a l o f th e n o n - t r e a t e d p l a n t s .

T h e s e d a ta c l e a r l y s h o w th a t p h o s p h o r u s

F i g u r e 12
P h o t o g r a p h (top) a n d a u t o r a d i o g r a m (b o tto m ) o f a n ’’a p e x to
a p e x " g r a f t u n io n b e tw e e n a l a t e r a l f r o m o n e t o m a t o p l a n t a n d th a t of
an o th er.

C a 45 w a s s u p p lie d to th e r o o t s of th e p l a n t f r o m w h ic h th e

l a t e r a l o r i g i n a t e d on th e le ft.

63

64

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65

c a n be a b s o r b e d by th e r o o t s y s t e m o f on e p la n t, t r a n s p o r t e d a c r o p e t a l l y
t h r o u g h i t s l a t e r a l , a c r o s s a g r a f t u n io n a n d b a s i p e t a l l y th r o u g h a l a t e r a l of
a s e c o n d p l a n t , a n d u t i l i z e d in th e d e v e l o p m e n t of i t s f r u i t .

G r a f t u n io n s in -

v o lv in g r e v e r s e d p o l a r i t y o f f e r l i t t l e r e s i s t a n c e to th e t r a n s p o r t of P

oo

.

R a d i o c a l c i u m , o n th e o t h e r h and, d o e s not r e a d i l y t r a n s g r e s s a g r a f t
u n io n f o r m e d by a p i c a l l y g r a f t i n g f o r c e d l a t e r a l s t o g e t h e r .

H ig h ly s i g n i f i ­

c a n t d i f f e r e n c e s o c c u r r e d b e tw e e n th e l a t e r a l s e g m e n t on th e t r e a t e d s id e
o f th e g r a f t u n io n a n d th e g r a f t u n ion s e g m e n t o r th e l a t e r a l s e g m e n t on th e
n o n - t r e a t e d s i d e of th e g r a f t u n io n .
p a r e n t by

12

A l a r g e a c c u m u l a t i o n of C a ^ ^ w a s a p ­

d a y s on th e s i d e of th e g r a f t un io n f o r m e d by th e p la n t s u p p lie d

th e is o t o p e a n d in th e g r a f t u n io n ( T a b le VII).

T h a t Ca

45

t r a n s p o r t w a s not

c o m p l e t e l y b l o c k e d i s s h o w n in F i g u r e 12 ( A u t o r a d i o g r a m ) .

No Ca

45

was

r e c o v e r e d in th e i m m a t u r e f r u i t s on th e p l a n t not r e c e i v i n g th e is o to p e .
In a s e c o n d s e r i e s th e s t e m of one t o m a t o p l a n t w a s i n a r c h e d in to
th e s t e m of a n o t h e r .

In th e d i s c u s s i o n w h ic h fo llo w s , th e s to c k w ill be

c o n s i d e r e d a s th e p l a n t b e in g g r a f t e d , a n d th e s c io n a s th e s e c o n d p la n t,
w h ic h s u p p l i e d th e top ( s c io n ) , h o w e v e r , a l s o r e t a i n e d its own r o o t s y s t e m .
T h u s , p r e s u m a b l y th e s c i o n h a d th e b e n e f it o f i t s own r o o t s a s w e ll a s th o s e
o f th e s t o c k ( F i g u r e 13).
g r o u p s , A a n d B.

T h e i n a r c h e d p l a n t s w e r e s e p a r a t e d into two

G r o u p A w a s t r e a t e d by a d d in g 100 m i c r o c u r i e s of Ca

to th e s o i l in w h ic h th e s t o c k p l a n t (a) w a s g ro w in g .
th e s c i o n p l a n t (b) a b s o r b e d th e i s o t o p e .

45

In g r o u p B th e r o o t s of

P l a n t s w e r e h a r v e s t e d 48 h o u r s

F i g u r e 13
P h o t o g r a p h s (left) a n d a u t o r a d i o g r a m s ( r ig h t) of i n a r c h e d s t e m s of
t o m a t o p l a n t s s h o w in g th e d i s t r i b u t i o n of r o o t a b s o r b e d r a d i o c a l c i u m
( C a ^ ) 48 h o u r s a f t e r t r e a t m e n t .
(T o p ) R a d i o c a l c i u m a p p l i e d to th e s o il in w h ic h th e s t o c k p la n t
(a) w a s g ro w in g .
(B ottom ) R a d i o c a l c i u m a p p l i e d to th e s o il in w h ic h th e s c i o n p la n t
(b) w a s g ro w in g .

66

67

a fte r Ca

w a s a d d e d to th e r e s p e c t i v e r o o t c u l t u r e s a n d a l l l e a v e s a n d s i d e -

s h o o ts w e r e r e m o v e d f r o m th e s t e m s .

T h e s t e m s w e r e th e n p r e p a r e d f o r

a u t o r a d i o g r a p h y a n d e x p o s e d to K o d a k X - r a y film (Blue B ra n d ).
A s w ith a p i c a l l y g r a f t e d l a t e r a l s ,

r e s u l t s w ith i n a r c h e d t o m a to

p l a n t s s u g g e s t e d l i t t l e o r no b a s i p e t a l t r a n s p o r t of C a 4 5 ( F i g u r e 13).
Ca

45

W hen

w a s s u p p l i e d to th e s t o c k r o o t s y s t e m th e c a l c i u m f o llo w e d th e t r a n s ­

p i r a t i o n s t r e a m t h r o u g h th e a p p r o a c h g r a f t a n d in to th e s c io n .

T h e re w as

no e v i d e n c e o f b a s i p e t a l t r a n s p o r t in th e s t e m of th e s c i o n p la n t, ( F i g u r e
13, T o p ).

L ik e w ise , w hen Ca

45

w a s a p p l i e d to th e r o o t s y s t e m of th e s c io n

p l a n t ( F i g u r e 13, B o tto m ), th e c a l c i u m m o v e d up th e v a s c u l a r s y s t e m of th e
s c io n a n d v e r y l i t t l e w a s o b s e r v e d in th e s to c k .

E x p e r i m e n t 11

T o f u r t h e r e v a l u a t e t h e e f f e c t o f g r a f t u n io n s on C a

45

tr a n s p o r t and

to s e p a r a t e th e e f f e c t s o f g r o w t h a n d s c i o n o r i e n t a t i o n v i g o r o u s l y g r o w in g
t o m a t o p l a n t s w e r e p r e p a r e d ( s e e F i g u r e 14) a s fo llo w s :
(a) A t w o - i n c h s t e m s e g m e n t c o n ta i n in g one bud w a s g r a f t e d
in th e u p r i g h t p o s i t i o n on th e d e c a p i t a t e d s te m of th e s a m e p la n t.
(b) A t w o - i n c h s t e m s e g m e n t c o n t a i n i n g one b u d w a s g r a f t e d
in a n i n v e r t e d p o s i t i o n a s in (a) a b o v e .
(c) P l a n t s w e r e d e c a p i t a t e d a t th e a p p r o x i m a t e h e ig h t of

F i g u r e 14
T op : P h o to g r a p h s h o w in g g r o w th of s c i o n s on d e c a p i t a t e d to m a to
p l a n t s 21 d a y s a f t e r b u d d in g .

(L e ft) bu d i n v e r t e d , ( c e n t e r ) bu d in n o r m a l

p o s i t i o n , a n d ( r ig h t) c o n t r o l (not b u d d e d ).
B ottom : P h o to g r a p h s h o w in g g r o w th of s c io n on g r a f t e d to m a to
p l a n t s 11 d a y s a f t e r g r a f t i n g .

(L eft) s c io n i n v e r t e d ,

n o r m a l p o s i t i o n , a n d (r ig h t) c o n t r o l (not g r a f t e d ) .

( c e n t e r ) s c i o n in

68

69

th e s c i o n in (a) a n d (b) a b o v e .

(No g r a f t un ion ).

(d) A s h i e l d w ith o n e bu d w a s b u d d e d to th e s t o c k in th e u p ­
r i g h t p o s i t i o n a n d t h e s t o c k d e c a p i t a t e d a b o v e the s h ie l d .
(e) A s h i e l d w ith o n e b u d w a s b u d d e d in a n i n v e r t e d p o s i t i o n
a n d th e s t o c k w a s d e c a p i t a t e d a b o v e th e s h ie l d .
(f) P l a n t s w e r e d e c a p i t a t e d a t h e ig h ts c o m p a r a b l e to th e b u d ­
d e d p l a n t s (d) a n d (e) a b o v e .
In a n a t t e m p t to r e d u c e p l a n t v a r i a b i l i t y in th e a b o v e s e r i e s , th e s c io n a n d
s t o c k w e r e o f th e s a m e c lo n e .
U po n c o m p l e t i o n o f g r a f t i n g o r b u d d in g th e p l a n t s w e r e p l a c e d in a
h u m id ity c h a m b e r (9 5 -1 0 0 % R. H . ) f o r fo u r d a y s a f t e r w h ic h th e y w e r e g ro w n
u n d e r lo n g d a y s o f e a r l y s u m m e r w ith f r e q u e n t f e r t i l i z a t i o n (w e e k ly a p p l i c a ­
tio n s of 100 m l o f 1 0 - 5 2 - 1 7 ) .
T e r m i n a l g r o w t h o f th e g r a f t e d s c i o n s w e r e r e c o r d e d 0, 11 a n d 24
days a f te r g ra ftin g .

E i g h t e e n d a y s a f t e r g r a f t i n g a n d 21 d a y s a f t e r budding,

w h en th e v a r i o u s u n io n s w e r e w e ll f o r m e d , 20 m i l l i l i t e r s of a C a
tio n ( 2 .

5

45

so lu ­

u c / m l ) w a s a p p l i e d to th e s o il in w h ic h e a c h p l a n t w a s g ro w in g .

T w o a n d s i x d a y s a f t e r th e p l a n t s w e r e s u p p l i e d w ith C a ^ th e y w e r e h a r ­
v e ste d .

T h e s c i o n s a n d g r a f t u n io n s in g r a f t e d p l a n t s a n d th e s c i o n s a n d

b ud u n io n s in b u d d e d p l a n t s w e r e s e p a r a t e d a n d d r i e d in a f o r c e d a i r o v e n at
7 0 ° C.

D r y w e i g h t s w e r e d e t e r m i n e d a n d th e t i s s u e a s s a y e d f o r r a d i o c a l c i u m .

70

R a d i o a c t i v i t y w a s e x p r e s s e d a s c o u n t s p e r m i n u t e p e r g r a m d r y w e ig h t o r
to ta l co u n ts p e r m in u te p e r p la n t p a r t.

A u t o r a d i o g r a m s w e r e p r e p a r e d of

s o m e o f t h e t r e a t e d p l a n t s a n d a r e i l l u s t r a t e d in F i g u r e 15.
T e r m i n a l g r o w t h a n d d r y w e ig h ts o f th e s c i o n s a r e r e c o r d e d in
T a b l e V III.

F r o m t h e s e d a ta it is a p p a r e n t t h a t th e i n v e r t e d s c io n w a s slo w

to i n i t i a t e g r o w th , a s c o m p a r e d to th e n o r m a l l y g r a f t e d s c io n o r th e c o n t r o l .
T h i s m a y i n d i c a t e t h a t a l o n g e r p e r i o d o f t i m e w a s r e q u i r e d f o r an a d e q u a te
u n io n to f o r m , o r t h a t by i n v e r s i o n s o m e e s s e n t i a l f a c t o r f o r g r o w th w a s
t e m p o r a r i l y e x c l u d e d f r o m th e s c io n .

It a p p e a r s ( T a b l e VIII, B ottom ) th a t

th e s lo w f o r m a t i o n of th e u n io n w a s th e c a u s a t i v e f a c t o r .

T h e s c i o n s on

th e n o n - i n v e r t e d g r a f t e d a n d c o n t r o l p l a n t s e l o n g a t e d 5. 5 to
in

11

d a y s , w h e r e a s th e i n v e r t e d s c i o n e l o n g a t e d o nly

0 .6

6

. 7 c e n tim e te rs

c en tim e te rs.

H o w e v e r , f r o m t h e 11th to th e 2 4 th d a y b o th s c i o n s a n d c o n t r o l t r i p p l e d
th e ir h e ig h ts.

T h e r a t e o f g r o w t h f r o m th e 11th to th e 2 4 th d a y o f th e i n v e r t e d

s c i o n a p p r o a c h e d t h a t of th e n o n - i n v e r t e d s c i o n a n d c o n t r o l .

T h e r e w e r e no

s i g n i f i c a n t d i f f e r e n c e s in d r y w e ig h t o r t e r m i n a l g r o w th of th e n o r m a l l y
g r a f t e d s c i o n s a n d th e c o n t r o l s ( T a b le VIII, T o p ).

T h e r e d u c t i o n s in d r y

w e ig h t a n d t e r m i n a l g r o w t h of th e i n v e r t e d s c i o n s w e r e h ig h ly s ig n if ic a n t
w h e n c o m p a r e d to th e n o r m a l l y g r a f t e d o r n o n - g r a f t e d ( c o n tr o l) p l a n t s .
I n v e r t i n g a b u d d id n o t a f f e c t th e s c i o n g r o w t h a s d r a s t i c a l l y a s i n ­
v e r t i n g t h e s c i o n ( F i g u r e 14).

T h e d r y w e ig h t o f th e s c i o n of a n i n v e r t e d

F i g u r e 15
A u t o r a d i o g r a m s i l l u s t r a t i n g th e d i s t r i b u t i o n of r a d i o c a l c i u m a s
i n f l u e n c e d by s c io n o r i e n t a t i o n in b u d d e d a n d g r a f t e d t o m a t o p l a n t s 48
h o u r s fo llo w in g a p p l i c a t i o n of Ca

45

to th e r o o t m e d i u m .

T op : B udded p l a n t s , (le ft) b u d not i n v e r t e d ; ( r i g h t ) bu d i n v e r te d .
Bottom: G r a f t e d p l a n t s , (le ft) s c i o n not i n v e r t e d ; ( r i g h t ) s c io n
in v e rte d .

71

TA BLE VIII
G R O W T H O F SC IO N ( T O M A T O /T O M A T O ) AS IN F L U E N C E D BY O R IE N T A ­
T IO N O F SCION
D ays A fte r
G ra ftin g

T r e a t m e n t of S c io n
In v e rte d

N ot I n v e r t e d

C o n tro l
(Not G r a f t e d )

( G r a m s d r y w e ig h t - m e a n s of
20

0

.

24

0

M eans

6

re p lic atio n s)

2. 2 0

2. 42

1. 55

. 23

4. 49

5. 01

3. 24

0. 13

3. 35

3. 72

02

L . S. D. t r e a t m e n t : 1 %

0. 67

L. S. D. t i m e :

0. 54

1

M ean
F o r T im e

%

( L e n g th of t e r m in a l g r o w th in c e n t i m e t e r s )
0

(in itial
le n g th )

1. 5

1

.

6

.5

1

8

.

5.

11

2

.

1

7. 1

24

6

.

6

24. 2

28. 3

.

12. 7

3. 4

M eans

1 1

0

L. S. D. t r e a t m e n t : 1%

2. 3

L. S. D. t i m e :

1. 9

1

%

.5

1

2

8

19. 7

73

b u d w a s o n ly o n e - h a l f o f t h e n o r m a l l y b u d d e d s c i o n a n d o n e - t h i r d o f th e
c o n t r o l ( T a b l e IX).

A s t i m e p r o g r e s s e d th e g r o w th r a t e of th e i n v e r t e d

b u d s a p p r o a c h e d t h a t of th e n o r m a l l y b u d d e d p l a n t s a n d n o n - b u d d e d c o n t r o l s .
A n a l y s i s o f th e g r a f t u n io n f o r r a d i o c a l c i u m d e r i v e d f r o m r o o t a b ­
s o r p t i o n s h o w e d no s i g n i f i c a n t d i f f e r e n c e s a m o n g th e u n io n s f o r m e d by th e
i n v e r t e d s c io n , n o n - i n v e r t e d s c i o n o r a c o m p a r a b l e s e g m e n t on th e n o n ­
g r a f t e d p l a n t ( T a b l e s X a n d XI).

L i k e w i s e , no d i f f e r e n c e s o c c u r r e d in th e

b u d u n io n of th e b u d d e d p l a n t s ( T a b l e s XII a n d XIII).
L e s s r a d i o c a l c i u m ( to t a l c p m ) w a s r e c o v e r e d in th e i n v e r t e d s c io n
t h a n th e n o r m a l l y g r a f t e d o r th e s c i o n s of th e n o n - g r a f t e d p l a n t s ( T a b le X);
y e t th e g r a f t u n io n s of i n v e r t e d , n o n - i n v e r t e d a n d a c o m p a r a b l e s e g m e n t in
th e c o n t r o l p l a n t c o n t a i n e d a b o u t e q u a l q u a n t i t i e s of C a ^ .

T h i s w o u ld i n ­

d i c a t e t h a t C a ^ i s b e in g t r a n s p o r t e d to, bu t n o t a s r a p id ly , a c r o s s th e
g r a f t u n io n s o f t h e i n v e r t e d s c i o n s .
In o r d e r to f u ll y e v a l u a t e th e in f l u e n c e o f th e g r a f t u n io n in r e l a t i o n
to t r a n s p o r t , th e d a ta w e r e e x p r e s s e d in c o u n ts p e r m i n u t e p e r g r a m d r y
w e ig h t.

T h e q u a n t i t y of C a

45

( c p m / g m d r y w t . ) r e c o v e r e d in th e g r a f t u n io n s

o r s c i o n s d id n o t d i f f e r s i g n i f i c a n t l y a m o n g th e in v e r t e d , n o n - i n v e r t e d o r
c o n t r o l p l a n t s ( T a b l e XI).

T h i s s u g g e s t e d th a t th e r a t e of g r o w th w a s th e

d e t e r m i n i n g f a c t o r in c a l c i u m t r a n s p o r t a n d no t th e n a t u r e of th e g r a f t un io n.
P o l a r i t y o f t h e t i s s u e w a s s t i l l a n i m p o r t a n t f a c t o r in c a l c i u m t r a n s p o r t , fo r
a s s o o n a s g r o w t h c o m m e n c e d t h e p o l a r i t y w a s r e o r i e n t e d in th e s c io n .

74

T A B L E IX
G R O W T H O F SC IO N (T O M A T O /T O M A T O ) AS IN F L U E N C E D BY O R IE N T A
T IO N O F BUD

D ays A fter
B udding

I n v e r t e d Bud

N o n - I n v e r t e d Bud

C o n t r o l (Not Budded)

( g r a m s d r y w e ig h t - m e a n s of 4 r e p l i c a t i o n s )
23

0 .7 3

1 .8 2

2 .4 4

29

1 .9 9

2 .7 0

3 .9 9

1 .3 6

2 .2 6

3 .2 2

M ean

L . S . D. f o r d i f f e r e n c e s in bu d o r i e n t a t i o n : 5%

0 .7 5

L . S . D. f o r d i f f e r e n c e s in b u d o r i e n t a t i o n :

1 .0 4

1%

75
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S i m i l a r t r a n s p o r t t r e n d s w e r e i n d i c a t e d in b u d d e d p l a n t s .

R ad io ­

c a l c i u m r e c o v e r e d a s t o t a l c o u n t s in th e s c i o n w a s s i g n i f i c a n t l y l e s s f o r th e
i n v e r t e d b u d ( T a b l e XII).
( T a b l e XII).

H o w e v e r , th e b u d u n io n s d id n ot d i f f e r s i g n i f i c a n t l y

T h e q u a n t i t y of C a ^ 5 ( c p m / g m d r y w t . ) r e c o v e r e d in th e s c i o n s

a n d b u d u n io n s o f th e b u d d e d t o m a t o p l a n t s is r e c o r d e d in T a b l e XIII.

W hen

th e d a ta w e r e e x p r e s s e d a s c o u n ts p e r m i n u t e p e r g r a m d r y w e ig h t, no s i g ­
n i f i c a n t d i f f e r e n c e s w e r e o b s e r v e d in th e s c i o n s o r b u d u n io n s .

It a p p e a r s

t h a t g r o w t h o f t h e s c i o n a n d n o t th e n a t u r e of th e bud u n io n is th e p r i m a r y
f a c t o r in th e u p w a r d t r a n s p o r t o f C a

45

E x p e r i m e n t 12

In e a r l i e r e x p e r i m e n t s it w a s s h o w n t h a t C a
t h r o u g h a g r a f t u n io n in v o lv in g r e v e r s e d p o l a r i t y .
m o v e d a c r o s s s u c h a u n io n .

45

w a s not t r a n s p o r t e d

C o n v erse ly , P

32

re a d ily

It w a s f u r t h e r o b s e r v e d t h a t th e g r o w th r a t e

o f t h e s c i o n w a s a m o r e i m p o r t a n t f a c t o r in C a 4 5 t r a n s p o r t th a n w a s th e
n a t u r e of t h e g r a f t u n io n .

T h e f o llo w in g e x p e r i m e n t w a s d e s i g n e d to e v a l u a t e

th e e f f e c t o f p h l o e m p o l a r i t y upon th e b a s i p e t a l t r a n s p o r t of f o l i a r a p p lie d
r a d i o p h o s p h o r u s in t h e a p p le .
In A u g u s t o f 1954, 14 u n i f o r m M a i l i n g I a p p le t r e e s ( o n e - y e a r - o l d )
w e r e s e l e c t e d f r o m th e n u r s e r y a t th e U n i v e r s i t y f a r m .

Two o n e - c e n tim e te r

s e g m e n t s o f p h l o e m ( b a r k ) a p p r o x i m a t e l y tw o to t h r e e c e n t i m e t e r s a p a r t

80

a n d 15 c e n t i m e t e r s a b o v e th e g r o u n d l e v e l w e r e i n v e r t e d on s e v e n of th e t r e e s .
T h e v e r t i c l e i n c i s i o n s o f th e p h l o e m r i n g s w e r e p l a c e d o p p o s ite o n e a n o t h e r
a n d h e l d in p l a c e by r u b b e r g r a f t i n g b a n d s .
tre e s,

On th e s e v e n r e m a i n i n g M a i l i n g I

tw o o n e - c e n t i m e t e r p h lo e m s e g m e n t s w e r e r e m o v e d a n d r e p l a c e d

w ith o u t i n v e r t i n g , a n d w ith th e v e r t i c l e i n c i s i o n s o p p o s i t e o n e a n o t h e r .
In t h e f o llo w in g s p r i n g th e y o u n g t r e e s w e r e p o t t e d a n d m o v e d in to
th e g r e e n h o u s e .
m i c r o c u r i e s of P

T e r m i n a l g r o w th w a s r e c o r d e d .
82

A f t e r te n w e e k s ,

100

w e r e a p p l i e d to a s id e s h o o t a t a p p r o x i m a t e l y c o m p a r a b l e

d i s t a n c e s a b o v e th e u p p e r p h lo e m r in g .

P l a n t s w e r e h a r v e s t e d 48 h o u r s

a f t e r t r e a t m e n t by r e m o v i n g a f i v e - c e n t i m e t e r p h lo e m s e g m e n t i m m e d i a t e l y
a b o v e th e p h l o e m r i n g s

a n d a s e c o n d s e g m e n t c o n ta i n in g th e p h lo e m r i n g s .

T h e s a m p l e s w e r e th e n p r e p a r e d a n d a s s a y e d f o r r a d i o a c t i v i t y .
W ith in tw o w e e k s a f t e r p e r f o r m i n g th e p h lo e m o r i e n t a t i o n t r e a t ­
m e n ts,

it w a s o b s e r v e d t h a t m a n y a d v e n t i t i o u s b u d s b e g a n to g r o w be lo w th e

p h lo e m r i n g s in th e n u r s e r y ; h o w e v e r , no s ig n i f i c a n t d i f f e r e n c e s w e r e o b ­
served.

In th e g r e e n h o u s e , th e fo llo w in g s p r i n g , th e n u m b e r of new s h o o ts

a r i s i n g on t r e e s b e lo w th e i n v e r t e d p h lo e m r i n g s w a s 70 p e r c e n t g r e a t e r
( T a b le XIV).

S i m i l a r r e s u l t s w e r e d e m o n s t r a t e d on to b a c c o w ith in fo u r

w e e k s a f t e r p h l o e m o r i e n t a t i o n t r e a t m e n t s ( T a b le XIV).

T h e t o ta l new top

g r o w th w h ic h o c c u r r e d on M a i l i n g I t r e e s , h a v in g tw o i n v e r t e d p h lo e m
r i n g s , w a s o n ly 50 p e r c e n t of t h a t w h ic h o c c u r r e d on the t r e e s not h a v in g
th e p h l o e m i n v e r t e d ( T a b l e XV).

T h i s a g r e e s w ith th e w o r k of S a x (1954)

81

T A B L E XIV
N U M B ER O F BUDS F O R C E D BELOW TW O P H L O E M RINGS ON A P P L E AND
T O B A C C O P L A N T S AS A R E S U L T O F R E M O V IN G , IN V E R T IN G AND
R E P L A C IN G , R E M O V IN G AND N O T R E P L A C IN G , OR R E M O V ­
ING AND R E P L A C IN G IN T H E N O R M A L PO SITIO N TW O
RIN G S O F P H L O E M

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83

w h e r e h e s u g g e s t e d th e p o s s i b i l i t y o f d w a r f i n g v i g o r o u s t r e e s by i n v e r t i n g
p h lo e m r i n g s .

F u r t h e r m o r e , a f o u r fo ld i n c r e a s e in th e s i z e o f th e s w e lli n g

w a s o b s e r v e d a b o v e th e u p p e r i n v e r t e d p h l o e m r i n g ( T a b le XV) c o m p a r e d
to n o n - i n v e r t e d p h l o e m r i n g s .
T h e e f f e c t o f t h e p h lo e m r i n g s on d o w n w a r d t r a n s p o r t of P ^ a r e
sh o w n in F i g u r e 16.

T h e q u a n t i t y of

(c p m /g m d ry w t.) re c o v e re d

a b o v e t h e i n v e r t e d p h l o e m r i n g s w a s g r e a t e r th a n in a c o m p a r a b l e s e g m e n t
a b o v e th e n o n - i n v e r t e d p h l o e m r i n g s .
p o rt of P

32

T h is s u g g e ste d th a t b a sip e ta l tr a n s -

w a s r e d u c e d by i n v e r t e d p h l o e m .

C o n s id e ra b le a m o u n ts of P

32

p a s s e d t h r o u g h th e u p p e r s e g m e n t o f i n v e r t e d p h lo e m ( F i g u r e 17, Bottom ),
p a r t i c u l a r l y i n t h e r e g i o n o f th e v e r t i c l e i n c i s i o n w h e r e th e x y le m a n d
p h lo e m w e r e r e g e n e r a t e d m o s t r a p i d l y .
p o r t r a y t h e d i s t r i b u t i o n of P
th e s t e m .

32

T h e a u t o r a d i o g r a m s ( F i g u r e 17)

in o n ly o n e p l a n e t h r o u g h a g iv e n r e g i o n of

If o n e c l o s e l y o b s e r v e s th e p a t t e r n in w h ic h th e n ew x y le m w a s

f o r m e d , th e r e g e n e r a t i o n w h ic h to o k p l a c e a t th e v e r t i c l e i n c i s i o n of th e
top p h l o e m r i n g f o ll o w e d a r o u n d to th e o p p o s i t e s id e of th e s t e m to th e
v e r t i c l e i n c i s i o n o f th e l o w e r p h l o e m r i n g ( s e e p h o to g r a p h , F i g u r e 16,
B o tto m ).

T h i s s u g g e s t e d th e d o w n w a r d t r a n s p o r t o f n u t r i e n t s w a s c h a n n e l e d

th r o u g h t h e v e r t i c l e i n c i s i o n s .

S u ch w o u ld a c c o u n t f o r th e a c c u m u l a t i o n

o f P ^ b e lo w th e s e c o n d p h l o e m i n v e r s i o n .

F i g u r e 16
Top: T h e c o m p a r a t i v e a m o u n t of f o l i a r a p p l i e d r a d i o p h o s p h o r u s
( c p m / g m d r y wt. ) r e c o v e r e d in th e v a r i o u s p h l o e m s e g m e n t s in th e ap p le
(E M I).

(A) F i v e c m s e g m e n t i m m e d i a t e l y a b o v e th e u p p e r i n v e r s io n ,

(B) five c m s e g m e n t c o n ta i n in g th e i n v e r t e d r i n g s , (C) fiv e c m s e g m e n t
i m m e d i a t e l y a b o v e th e u p p e r n o n - i n v e r t e d p h lo e m r in g , a n d (D) fiv e cm
seg m e n t co n tain in g n o n -in v e rte d rin g s.
Bottom: P h o to g r a p h s s h o w in g a s e g m e n t of th e a p p le s te m w ith
th e p h lo e m r e m o v e d i l l u s t r a t i n g x y le m r e g e n e r a t i o n in v i c i n i t y of p h lo e m
rin g s.

(L e ft) P h lo e m r i n g s w e r e i n v e r t e d .

N ote: S w e llin g a n d r e g e n e r a ­

tio n of x y le m a t s i t e of v e r t i c l e i n c i s i o n of i n v e r t e d p h l o e m r i n g s .
(R ight) P h lo e m r i n g s w e r e not i n v e r t e d .

CPM/GM

DRY WT

2000

1500

1000

500

0

85

86

S e c tio n C

S o m e F a c t o r s A f f e c t i n g T r a n s p o r t in th e P h lo e m

T h e l a c k o f b a s i p e t a l t r a n s p o r t of c a l c i u m in c o tto n h a s b e e n a t t r i ­
b u t e d to i t s i n a b i l i t y to m o v e in th e p h lo e m (M a s o n a n d M a s k e l l 1931).
P r e v i o u s e x p e r i m e n t s (10,

11

) on c a l c i u m m o b i l i t y s u g g e s t e d th e p o s s i b l e

r o l e o f p o l a r i t y in c o n t r o l l i n g th e t r a n s p o r t of c a l c i u m .

W ent a n d T h i m a n n

(1937) r e p o r t e d t h a t p o l a r i t y w a s t e m p o r a r i l y s u s p e n d e d by low c o n c e n t r a ­
tio n s of e t h e r v a p o r s .

A c c o r d i n g ly , th e fo llo w in g e x p e r i m e n t w a s d e s ig n e d

to a s c e r t a i n th e r o l e o f p o l a r i t y in c a l c i u m t r a n s p o r t .

E x p e r i m e n t 13
T h e e f f e c t o f d i - e t h y l e t h e r u po n th e t r a n s p o r t of f o l i a r a p p lie d
r a d io c a lc iu m w a s stu d ie d .

O ne o f th e fu lly e x p a n d e d p r i m a r y l e a v e s of th e

b e a n p l a n t ( v a r . B la c k V a le n ti n e ) w a s t r e a t e d by a p p ly in g f o u r m i c r o c u r i e s
of C a 4 5 s o l u t i o n a s a d r o p on th e m i d r i b .

P l a n t s w e r e th e n g r o u p e d a s

fo llo w s :
45
(a) T h e p r i m a r y l e a f t r e a t e d w ith C a
w as ex p o sed
f o r 96 h o u r s in a 500 m i l l i l i t e r g l a s s j a r a t r o o m
t e m p e r a t u r e to v a p o r s f r o m 25 m i l l i l i t e r s of e t h e r
(d i-eth y l).
(b) T h e l e a f o p p o s i t e th e C a
to e t h e r a s in (a) a b o v e .

45

,
tr e a te d le af w as exposed

87

(c) T h e C a 4 ^ t r e a t e d l e a f w a s p l a c e d in a c o m p a r a b l e
g l a s s j a r w ith no e t h e r .
(d) T h e l e a f o p p o s i t e th e C a 4 "* t r e a t e d l e a f w a s p l a c e d
in a c o m p a r a b l e g l a s s j a r w ith no e t h e r .
P l a n t s w e r e h a r v e s t e d 96 h o u r s a f t e r t r e a t m e n t w ith C a 4 ^ a n d a u t o r a d i o g r a m s
p rep ared .
W h e n th e C a
tra n s p o rt of Ca

45

p l a n t ( F i g u r e 18).

45

t r e a t e d l e a f w a s s u b j e c t e d to e t h e r c o n s i d e r a b l e b a s i p e t a l

o c c u r r e d o ut of th e t r e a t e d l e a f a n d in to o t h e r p a r t s of t h e
W h en th e l e a f o p p o s i t e th e C a

45

to e t h e r , l i m i t e d t r a n s p o r t o c c u r r e d w ith in th e C a

t r e a t e d leaf w as su b jec te d

45

t r e a t e d le a f .

No b a s i p e t a l

t r a n s p o r t o c c u r r e d in . he l e a v e s o f g r o u p s (c) o r (d) a b o v e .
S i m i l a r r e s u l t s w e r e o b t a i n e d w ith r a d i s h p l a n t s (v a r. Ic ic le ) s u b j e c t e d
to t r e a t m e n t s (a) a n d (c) d e s c r i b e d a b o v e .

W h en th e C a 4 ^ t r e a t e d l e a f w a s e x ­

p o s e d to e t h e r v a p o r s , c o n s i d e r a b l e b a s i p e t a l t r a n s p o r t o c c u r r e d .

P h o to g r a p h s

a n d a u t o r a d i o g r a m s ( F i g u r e 19) of c r o s s a n d lo n g itu d in a l s e c t i o n s of th e s w o lle n
h y p o c o ty l of r a d i s h s u g g e s t e d t h a t C a 45 w a s t r a n s p o r t e d b a s i p e t a l l y .

In t r e a t ­

m e n t (c) w h e r e th e C a 45 t r e a t e d l e a f w a s n o t s u b j e c t e d to e t h e r v a p o r , no b a s i p e tal tra n s p o r t of Ca

45

occurred.

U n d e r th e c o n d itio n s d e s c r i b e d a b o v e no

v i s i b l e i n j u r y f r o m a n e s t h e t i z a t i o n o c c u r r e d to th e t r e a t e d p l a n t s .

88

89

E x p e r i m e n t 14
T h e a b s o r p tio n a n d su b se q u e n t tr a n s p o r t of fo lia r a p p lied c h e la te d
r a d i o c a l c i u m w a s s t u d i e d in b e a n p l a n t s .

R a d i o c a l c i u m (408 uc) w a s a d d e d

to t e n m i l l i l i t e r s of a t e n p e r c e n t s o lu tio n of d i - s o d i u m e t h y l e n e d i a m i n e t e t r a ­
a c e ta te

a n d s a t u r a t e d s o l u t i o n s of p i c r o l i n i c a c id , 2, 5 - d i c h l o r o - 3 , 6 - d i h y d r o x y -

p -q u in o n e , an d l- a m in o - 2 - n a p t h o l- 4 - s u lf o n ic a c id .
w a s a d j u s t e d to 8. 0, 1 . 5 ,

1 . 5 a n d 2. 0 r e s p e c t i v e l y w ith p o t a s s i u m h y d r o x id e

o r h y d r o c h l o r i c a c i d a n d a e r a t e d f o r 24 h o u r s .
of 408 m i c r o c u r i e s of C a

T h e pH of th e s o lu tio n s

45

T h e c o n t r o l s o lu tio n c o n s i s t e d

C l 2 in te n m i l l i l i t e r s of d i s t i l l e d w a t e r w ith th e pH

a d j u s t e d to 2. 0 w ith h y d r o c h l o r i c a c i d .

A f t e r 24 h o u r s a o n e - m i l l i l i t e r a liq u o t

of e a c h of th e a b o v e f o u r c h e l a t e d p r e p a r a t i o n s w a s r e m o v e d , and an e x c e s s
o f 2N o x a l i c a c i d a d d e d .
c a t e d c h e l a t i o n of t h e C a

T h e a b s e n c e of a p r e c i p i t a t e ( c a lc iu m o x a la te ) in d i45

T h e c h e l a t e d r a d i o c a l c i u m (0. 01 m l) w a s a p p l i e d to th e m i d r i b of a
f u lly e x p a n d e d p r i m a r y l e a f o f th e b e a n ( v a r . B ountiful).

A f t e r 24 h o u r s p l a n t s

w e r e h a r v e s t e d a n d s e p a r a t e d a s fo llo w s : (a) t r e a t e d le a f, (b) s te m p l u s n o n ­
t r e a t e d le a f , a n d (c) r o o t s .

O th e r p la n ts w e re p r e p a r e d fo r au to ra d io g ra p h y .

P i c r o l i n i c a n d l - a m i n o - 2 - n a p t h o l - 4 - s u l f o n i c a c i d s r e s u l t e d in s e v e r e
b u r n i n g o f th e t r e a t e d le a f .
c h ela tio n .

T h i s li k e l y r e s u l t e d f r o m th e low pH r e q u i r e d f o r

R a d i o a c t i v e a s s a y o f th e v a r i o u s p l a n t p a r t s r e v e a l e d th a t no C a

w a s t r a n s p o r t e d o u t of th e t r e a t e d l e a v e s i r r e s p e c t i v e of c h e la tio n .
w e r e s u b s t a n t i a t e d by a u t o r a d i o g r a m s .

45

The r e s u lts

A ll s o l u t i o n s e n t e r e d th e l e a v e s but w e r e

t r a n s p o r t e d o n ly t o w a r d th e p e r i p h e r y .
*R e c r y s t a l i z e d d i - s o d i u m e t h y l e n e d i a m i n e t e t r a a c e t a t e w a s s u p p lie d by th e
A l r o s e C h e m i c a l C o . , P r o v i d e n c e , R- I.

91

V . D ISC U SSIO N

S to ck -S cio n In te rre la tio n s h ip

W ith t h e a d v e n t o f r a d i o i s o t o p e s a n d r e l a t e d te c h n i q u e s it h a s b e e n
p o s s i b l e w i t h c e r t a i n h o r t i c u l t u r a l p l a n t s to s tu d y th e r o l e o f th e r o o t s t o c k ,
s c i o n a n d g r a f t u n io n in a b s o r p t i o n a n d s u b s e q u e n t t r a n s p o r t of n u t r i e n t s .
Both q u a n t i t a t i v e a n d v i s u a l r e s u l t s h a v e b e e n o b ta in e d f o r t h e u p w a r d a n d
dow nw ard tr a n s p o r t of n u trie n ts .

T h e s e s t u d i e s h a v e b e e n c o n c e r n e d c h ie f ly

w ith p h o s p h o r u s a n d c a l c i u m ; p h o s p h o r u s , b e c a u s e of i t s i m p o r t a n c e in th e
m e t a b o l i c p r o c e s s e s o f th e p la n t, a n d c a l c i u m ,

b e c a u s e of r e s t r i c t i o n of

g ro w th w hen a b se n t.
It i s b e l i e v e d t h a t t h e c r i t i c a l t i m e in t h e g r o w th o f th e s c i o n on an y
r o o t s t o c k i s in th e s p r i n g w h e n g r o w t h i s r a p i d a n d th e m e t a b o l i c p r o c e s s e s
of th e p la n t a r e a t a m a x im u m .

If a t t h i s t i m e , f o r s o m e r e a s o n , th e r o o t -

s t o c k c a n n o t s u p p ly th e n e c e s s a r y m e t a b o l i t e s , it i s c o n c e i v a b l e t h a t s c io n

v
g r o w t h w ill b e a f f e c t e d .

F o r th e s e r e a s o n s e x p e r im e n ts re p o rte d , h erein ,

w e r e o f r e l a t i v e l y s h o r t d u r a t i o n a n d d u r i n g p e r i o d s of h ig h m e t a b o l i c a c ti v ity .
It w a s s h o w n t h a t r o o t e d c u t t i n g s o f M a i l i n g (EM) VII, IX, a n d XVI
h a v e d i f f e r e n t c a p a c i t i e s to a b s o r b r a d i o p h o s p h o r u s (P
ti o n s .

32

) fro m n u trien t s o lu ­

A b s o r p t i o n w a s c l o s e l y r e l a t e d to th e v i g o r o b s e r v e d in t h e s e s t o c k s

u n d e r fie ld c o n d itio n s.

T he r e s u lts p re s e n te d h e re for P

32

a b so rp tio n a g re e ,

92

in p a r t , w i t h t h o s e of P e a r s e (1939) f o r n i t r o g e n a n d p o t a s s i u m .

P earse

sh o w ed a d if f e r e n tia l a b s o r p tio n of p o ta s s iu m fo r d w a rfin g and v ig o ro u s
r o o t s t o c k s e a r l y in th e s e a s o n , b u t a s th e s e a s o n p r o g r e s s e d t h i s d i f f e r e n c e
d isa p p e a re d .

B e r r y (1938) m e a s u r e d t h e a b s o r p t i o n o f p o t a s s i u m b r o m i d e

a n d c o n c l u d e d t h a t r o o t s t o c k s d i f f e r e d in t h e i r a b s o r p t i o n c a p a c i t i e s .

He

fo u n d th e g r e a t e s t a b s o r p t i o n w ith E M XII, E M II w a s i n t e r m e d i a t e , w ith
t h e l o w e s t v a l u e s f o r E M IX.

S w a r b r i c k e t a l (1946) s u g g e s t e d t h a t th e

d w a r f i n g e f f e c t m a y b e t h e r e s u l t o f a d i f f e r e n t i a l r a t e of a b s o r p t i o n o r t r a n s lo c a t i o n , o r a c o m b i n a t i o n o f th e tw o .
In a s i m i l a r m a n n e r , p h lo e m t r a n s p o r t of f o l i a r a p p l i e d P

32

in

M c I n t o s h s c i o n s on E M VII, IX a n d XVI a n d D e lic i o u s s e e d l i n g r o o t s t o c k s
v e r y c l o s e l y p a r a l l e l th e s c i o n v i g o r o b s e r v e d on t h e s e s t o c k s u n d e r f ie ld
c o n d itio n s.
th a t P

32

No m a r k e d a c c u m u l a t i o n o c c u r r e d a t th e b u d u n io n s i n d i c a t i n g

p a s s e d f r e e l y t h r o u g h th e p h lo e m p o r t i o n of th e g r a f t .

T h e re w as

no s i g n i f i c a n t a c c u m u l a t i o n a b o v e th e b u d u n io n s .
It a p p e a r e d t h e n t h a t t h e s c i o n (M c In to s h ) w a s s o m e h o w m o d if ie d ,
e i t h e r t h r o u g h i t s a b s o r p t i o n o r t r a n s p o r t m e c h a n i s m , by th e s to c k .

That

r e d u c e d t r a n s p o r t i n t h e s t o c k of E M IX c a n a c c o u n t f o r t h e l e s s e r q u a n tity
o f P 32 r e c o v e r e d

f r o m th e f o l i a r a p p l i c a t i o n in th e r o o t s i s n o t t e n a b l e .

If

t h i s b e th e c a s e , t h e n a m a r k e d i n c r e a s e o f P 32 w o u ld h a v e r e s u l t e d in th e
scio n .

T h e f o r m in w h ic h th e a p p l i e d p h o s p h o r u s w a s t r a n s p o r t e d h a s no t b e e n

e sta b lish e d .

93

In c o n t r a s t to p h o s p h o r u s , C a

45

a p p l i e d to a p p le f o lia g e f a i l e d to

m o v e b a s i p e t a l l y f r o m th e s i t e of a p p l i c a t i o n .
In r o o t a b s o r p t i o n s t u d i e s of P 32 a n d C a 43 u s i n g s c i o n s o f M c I n to s h
on E M VII, IX a n d XV I a n d D e l i c i o u s s e e d l i n g r o o t s t o c k s c o n s i s t e n t d i f f e r ­
ences w ere observed.

T h e m o s t d w a r f i n g s t o c k s (E M IX a n d VII) a b s o r b e d

a n d t r a n s p o r t e d t h e l e a s t q u a n t i t y of b o th P 32 a n d C a 45 to th e s c i o n s b u d d e d
th e re o n .

V i g o r o u s s t o c k s s h o w e d i n c r e a s e d a b s o r p t i o n a n d t r a n s p o r t to th e

s c i o n o f b o th P
and Ca

45

32

and Ca

45

.

In g e n e r a l , a q u a l i t a t i v e c o m p a r i s o n o f th e P

qo

r e c o v e r e d in th e s c i o n s on E M VII, IX, a n d XVI o r D e lic i o u s s e e d ­

li n g w a s c l o s e l y r e l a t e d to v i g o r .
QO

A s w ith th e d o w n w a r d t r a n s p o r t of P ^ f r o m f o l i a r a p p lic a tio n , no
sig n ific a n t a c c u m u la tio n of P
ro o t a p p lic a tio n s.

32

o r Ca

45

w e r e o b s e r v e d a t th e b u d u n ion f r o m

T h i s i n d i c a t e d t h a t th e bu d union, p e r se, d id not s u b s t a n -

t i a l l y i n t e r f e r e w ith t h e u p w a r d o r d o w n w a r d t r a n s p o r t o f P 32 o r th e u p w a r d
t r a n s p o r t of C a

45

.

T h e l e s s e r q u a n tity o f P

32

and Ca

45

r e c o v e r e d in th e

s c i o n s o n d w a r f i n g s t o c k s c a n th e n be a t t r i b u t e d e i t h e r to r e d u c e d a b s o r p t i o n
o r t r a n s p o r t , o r a c o m b i n a t i o n of th e tw o.

In v ie w of th e d a ta on p h lo e m

t r a n s p o r t a n d a b s o r p t i o n w ith r o o t e d c u ttin g s , it is h ig h ly s u g g e s t i v e th a t
t h e m a j o r c o n t r i b u t i n g f a c t o r r e s p o n s i b l e f o r th e l e s s e r q u a n t i t i e s of P

32

o r C a 45 r e c o v e r e d in t h e M c I n to s h s c i o n s on d w a r f in g r o o t s t o c k s is th e
l o w e r a b s o r b i n g c a p a c i t y o f th e r o o t s t o c k .

T r a n s p o r t m a y a l s o be a c o n t r i ­

b u t i n g f a c t o r s i n c e th e a b s o r p t i o n a n d t r a n s p o r t p r o c e s s e s a r e in t e r d e p e n d e n t

94

upon one a n o th e r .

No d i s t i n c t i o n , h o w e v e r , w a s m a d e b e tw e e n th e tw o

p ro cesses.
In s t u d i e s o f f o l i a r a b s o r p t i o n a n d t r a n s p o r t u s i n g d w a r f a n d n o n -

q9
d w a r f t o m a t o p l a n t s , th e l e s s e r q u a n t i t y o f P

r e c o v e r e d in th e r o o t s o f th e

g e n e t i c a l l y d w a r f e d t o m a t o p l a n t s w a s d e f i n i t e l y a t t r i b u t e d to r e d u c e d t r a n s ­
p o rt.

D w a rf an d n o n -d w a rf p la n ts a b s o rb e d a p p ro x im a te ly equal q u an titie s

of fo lia r ap p lie d P

32

( T a b l e IV).

H ow ever, m o r e P

32

w a s r e c o v e r e d in th e

t o p s ( m i n u s th e t r e a t e d le a f ) of th e d w a r f th a n in a c o m p a r a b l e s a m p l e of
th e n o n -d w a rf p la n t.

A c c o r d i n g l y , l e s s P^2 w a s r e c o v e r e d in th e r o o t of

th e d w a r f t h a n n o n - d w a r f t o m a t o p l a n t .

E f f e c t o f th e G r a f t U n io n a n d S c io n O r i e n t a t i o n
on th e T r a n s p o r t of P

32

and Ca

45

In th e p r e c e e d i n g s t u d i e s w ith c o m p a t i b l e c o m b i n a t i o n s of a p p le
s c i o n s a n d r o o t s t o c k s , th e ^ r a f t u n io n d id n o t a p p e a r to r e s t r i c t c a lc iu m o r
\

p h o sp h o ru s tra n s p o rt.

S t u d i e s d e s i g n e d to m o r e c r i t i c a l l y e v a l u a t e th e r o l e

o f th e g r a f t u n io n i n d i c a t e d no a p p a r e n t r e s t r i c t i o n of p h o s p h o r u s t r a n s p o r t .
C o n s i d e r a b l e v a r i a t i o n , h o w e v e r , w a s e v id e n t fo llo w in g r o o t a b s o r p t i o n of
p^.

T h i s u n d o u b te d ly r e s u l t e d f r o m u n c o n t r o l l a b l e f a c t o r s e x i s t i n g in th e

s o i l m e d i a , p a r t i c u l a r l y p h o s p h o r u s fix a tio n .

A ls o , it i s d iff ic u lt to o b ta in

a c o n s t a n t s p e c i f i c a c t i v i t y w h e n th e r a d i o i s o t o p e w a s a d d e d to s o il c u l t u r e s .

95

A n i n v e s t i g a t i o n o f th e a c r o p e t a l t r a n s p o r t o f N a2 2 , P ^ ,

S ^ , C l^ ,

r^Q45
C a , lwm5
M n 2 ' 54 , uF ^e 5 5 " 59 , pC u 64 , „Z n 65 , „R,b8 6 , „S r 89 , M o 99 , a n d Ba 1 4 0 t h r o u g h
g r a f t u n i o n s w a s p e r f o r m e d by m e a n s o f a u t o r a d i o g r a p h y .
g r a f t u n io n , p e r s e , i m p e d e th e u p w a r d t r a n s p o r t .

In no c a s e d id th e

S i m i l a r l y , it m a y be a s ­

s u m e d t h a t i t o f f e r e d no r e s i s t a n c e to th e t r a n s p o r t of th e n o n - r a d i o a c t i v e
e le m e n ts.
In c o n t r a s t to n o r m a l l y g r a f t e d s c i o n s , g r a f t u n io n s w h ic h in v o lv e d
r e v e rs e d p o la rity se v e re ly red u ced Ca
tra n s p o rt of P

32

.

45

t r a n s p o r t a n d to a l e s s e r d e g r e e

W h en f o r c e d l a t e r a l s of v i g o r o u s to m a to p l a n t s w e r e g r a f t e d

t o g e t h e r t e r m i n a l l y , a u n io n w a s f o r m e d in v o lv in g th e a p e x e s o f tw o p l a n t s .
R a d i o c a l c i u m (C a

45

) a p p l i e d to th e r o o t s of on e o f th e tw o p l a n t s a c c u m u l a t e d

in th e g r a f t e d l a t e r a l o f th e t r e a t e d p l a n t .

R a d i o p h o s p h o r u s on th e o t h e r hand,

a c c u m u l a t e d to a l e s s e r d e g r e e a n d w a s e v e n t u a l l y d i s t r i b u t e d th r o u g h o u t b o th
p la n ts.

T h e f a c t t h a t c a l c i u m i s n o t m o b i l e in th e p h lo e m a s s u g g e s t e d by

M a s o n a n d M a s k e l l (1931) m a y a c c o u n t f o r th e r e d u c e d t r a n s p o r t a c r o s s a
g r a f t u n io n in v o lv in g r e v e r s e d p o l a r i t y .

T o be t r a n s l o c a t e d to a n a d j a c e n t

p l a n t a t t a c h e d by g r a f t i n g , C a 45 m u s t be t r a n s p o r t e d b a s i p e t a l l y th r o u g h
e i t h e r th e p h lo e m o r x y lem .

S in c e c a l c i u m is p r e s u m a b l y i m m o b i l e in th e

p h l o e m a n d l i t t l e e v i d e n c e e x i s t s f o r th e d o w n w a r d t r a n s p o r t o f io n s o r m o r e
c o m p l e x c o m p o u n d s in t h e x y le m , th e r e s u l t s r e p o r t e d h e r e i n a p p e a r c o n s i s t e n t .
T h e t e r m i n a t i o n o f th e t r a n s p i r a t i o n s t r e a m s of a p e x to a p e x g r a f t e d l a t e r a l s

96

c a n b e v i s u a l i z e d a t t h e g r a f t u n io n ; t h e r e b y e a c h l a t e r a l o b t a i n s i t s c a l c i u m
r e q u ir e m e n t fro m its p a r e n t p la n t.

It i s c l e a r l y e v id e n t t h a t d i f f e r e n t m e c h ­

a n is m s a r e o p e ra tiv e for calc iu m an d p h o sp h o ru s tr a n s p o r t u n d er th e se c o n ­
d itio n s.
T h e d a t a h e r e i n r e p o r t e d s u g g e s t e d th a t p o l a r i t y a s a p h y s i o - e l e c t r i c a l
p h e n o m en o n m ay , u n d e r c e r t a i n co n d itio n s, g o v e rn c a lc iu m t r a n s p o r t .

T h is

w a s f u r t h e r s u p p o r t e d by r e c e n t w o r k o f L o w e n h a u p t (1956), who, u s i n g
a q u a t i c p l a n t s ( P o l a m o g e t o n c r i s p u s L . ) r e p o r t e d t h a t c a l c i u m m o v e d in to
t h e a b a x i a l l e a f s u r f a c e w h e n in c o n t a c t w ith th e a q u e o u s m e d i a a n d o u t o f
th e a d a x i a l s u r f a c e w h e n th e l a t t e r w a s in c o n t a c t w ith th e a q u e o u s m e d i a .
E s s e n tia lly , th e d ir e c tio n of c a lc iu m m o v e m e n t u n d e r lig h te d c o n d itio n s w as
in to th e l o w e r s u r f a c e o f th e l e a f a n d o u t o f th e u p p e r s u r f a c e .

In t h e d a r k

th e d i r e c t i o n w a s r e v e r s e d .
W hen C a

45

t r e a t e d b e a n l e a v e s w e r e s u b j e c t e d to lo w c o n c e n t r a t i o n s

o f e t h e r ( d i - e t h y l ) v a p o r , th e f o l i a r a p p l i e d C a

45

w a s t r a n s p o r t e d ou t o f t h e

t r e a t e d l e a f a n d to a l l o t h e r p a r t s o f th e p l a n t ( F i g u r e 18).
o c c u r r e d w ith r a d i s h ( F i g u r e 19).

S im ila r re s u lts

T h is m ay have re s u lte d fro m a te m p o ra ry

s u s p e n s i o n of p o l a r i t y a s a r e s u l t of a n e s t h e t i z a t i o n , a s r e p o r t e d by ^A^ent a n d
T h i m a n n (1 9 3 7 ).

O n th e o t h e r h an d, a c h a n g e in c e l l m e m b r a n e p e r m e a b i l i t y

h a s not been c o m p le te ly e lim in a te d .

It w o u ld a p p e a r , h o w e v e r , t h a t u n d e r

t h e c o n d i t i o n s of l o c a l i z e d a n e s t h e s i a only, th e c e l l m e m b r a n e s in c l o s e

97

p r o x i m i t y w o u ld b e a f f e c t e d .

S till, l o c a l a n e s t h e s i a i s s u f f i c i e n t p r o v i d i n g

45
th e C a

tis s u e is-a n e sth e tiz e d .

It is c o n c e i v a b l e t h a t a s a r e s u l t of t r e a t ­

m e n t w ith e t h e r , a t r a n s l o c a t a b l e c o m p o u n d is f o r m e d , w h ic h is c a p a b l e of
a f f e c t i n g th e p e r m e a b i l i t y o f th e m e m b r a n e s t h r o u g h w h ic h it p a s s e d .

If

t h i s be th e c a s e , th e n o n e c o u ld e x p l a i n th e t r a n s p o r t of f o l i a r a p p l i e d C a ^ " \
in th e p r e s e n c e o f d i - e t h y l e t h e r , on th e b a s i s of c e l l m e m b r a n e p e r m e a b i l i t y .
T h e a u t h o r i s not, h o w e v e r , a w a r e of s u c h a c o m p o u n d .
W h e n a t o m a t o s c i o n i s i n v e r t e d on a s to c k , th e t o ta l r o o t a p p lie d
Ca

45

t r a n s p o r t e d to th e s c i o n w a s l e s s th a n in n o r m a l l y g r a f t e d o r n o n - g r a f t e d

p la n ts.

L i n e a r g r o w t h a n d d r y w e ig h t of th e i n v e r t e d s c i o n s w e r e a l s o l e s s .

If, h o w e v e r , th e C a 45 r e c o v e r e d in th e i n v e r t e d s c io n is e x p r e s s e d a s p e r
u n it of d r y w e ig h t ( c p m / g m d r y w t . ) t h e n no d i f f e r e n c e s w e r e o b s e r v e d .
T h i s s u g g e s t e d t h a t th e r a t e of g r o w th of th e s c io n w a s th e f a c t o r d e t e r m i n i n g
c a l c i u m t r a n s p o r t a n d n o t th e n a t u r e of th e g r a f t u n io n.
a c c u m u l a t i o n a t o r b e lo w th e u n io n s .

T h e f a c t th a t Ca

T h e r e w a s no m a r k e d

45

t r a n s g r e s s e d the

u n io n f o r m e d by a n i n v e r t e d s c i o n d o e s not m a k e th e p o l a r i t y h y p o th e s i s u n ­
te n a b le .

A l o n g e r t i m e w a s r e q u i r e d f o r th e g r a f t u n io n to f o r m b e tw e e n th e

i n v e r t e d s c i o n a n d th e s to c k , but o n c e it w a s e s t a b l i s h e d a n d g r o w th o f th e
s c i o n c o m m e n c e d , th e g r o w t h r a t e a p p r o a c h e d th a t of th e n o r m a l l y g r a f t e d
p la n t.

T h i s s u g g e s t e d t h a t th e p o l a r i t y of th e i n v e r t e d s c io n w a s r e - e s t a b ­

l i s h e d a s t h e n e w p a t h w a y s of t r a n s p o r t w e r e e s t a b l i s h e d .

C o n v e r s e ly , w hen

98

tw o l a t e r a l s w e r e g r a f t e d t o g e t h e r t e r m i n a l l y , t h e r e w a s no n e e d f o r r e - e s t a b lis h m e n t of th e t r a n s p o r t p a th w a y s.

Z i m m e r m a n a n d C o n n a r d (1934) s u g g e s t e d

t h a t e l a b o r a t e d f o o d s , a s w e ll a s s a l t s , m o v e d u p w a r d a n d d o w n w a r d t h r o u g h
ste m s.

T h e y c o n c l u d e d t h a t li v i n g c e l l s ta k e a n a c t i v e p a r t in t r a n s l o c a t i o n ,

a n d t h a t p r o t o p l a s m i c s t r e a m i n g s h o u ld h a v e th e s a m e e f f e c t w h e t h e r th e
d i r e c t i o n be b a c k w a r d o r f o r w a r d .

T h i s , h o w e v e r , w a s n o t th e c a s e a s

h e re in r e p o r te d fo r calc iu m .
T h e o r i e n t a t i o n of th e p h lo e m h a s a p r o n o u n c e d e f f e c t upon th e d o w n ­
w a r d t r a n s p o r t of m e t a b o l i c p r o d u c t s in th e a p p le .

Sax (1954) s u g g e s t e d t h a t

by i n v e r t i n g r i n g s o f p h l o e m th e t r e e c a n be d w a rfe d , p r e s u m a b l y by p r e ­
v e n t i n g th e m o v e m e n t of n u t r i e n t s a n d a u x in s th r o u g h th e i n v e r t e d p h lo e m
c e lls.
ity .

S a x (19 54 ) a n d Sax a n d D i c k s o n (1956) a t t r i b u t e d t h i s to p h lo e m p o l a r ­
H o w e v e r , no q u a n t i t a t i v e d a t a w e r e p r e s e n t e d .

In th e p r e s e n t e x p e r i ­

m e n t s , th e g r o w t h of th e to p w a s d e f i n i t e l y r e s t r i c t e d a s a r e s u l t of i n v e r t i n g
tw o o n e - c e n t i m e t e r p h l o e m r i n g s ( T a b le XIV).

T h a t th e i n v e r t e d p h lo e m

r i n g s i n t e r f e r e d w ith a u x in t r a n s p o r t c a n b e a s s u m e d a s a r e s u l t of th e g r o w th
o f th e l a t e n t b u d s b e lo w th e r i n g s .

O ne y e a r a f t e r r i n g i n g a d v e n ti tio u s b u d s

c o n t i n u e d to b r e a k o n t h o s e t r e e s w ith i n v e r t e d p h lo e m r i n g s .

T h is m ay

h a v e r e s u l t e d f r o m th e i n a b i l i t y of a u x in , w h ic h te n d s to s u p p r e s s l a t e r a l
b u d d e v e l o p m e n t , to p a s s a c r o s s th e i n v e r t e d p h l o e m r i n g s .
M o re fo lia r a p p lie d

a c c u m u l a t e d a b o v e th e i n v e r t e d th a n a b o v e

99

th e n o n - in v e r te d p h lo em rin g s .
w as not d e te rm in e d .

T h e n a t u r e of th e a c c u m u l a t i o n p r o d u c t

D i c k s o n a n d S a m u e l s (1956) r e p o r t e d s i m i l a r r e s u l t s

o n a s i n g l e t r e e , a n d c o n c l u d e d t h a t th e d w a r f i n g r e s p o n s e o b s e r v e d m a y
have r e s u lte d fro m in p ed ed tr a n s p o r t of o rg a n ic n u trie n ts .
b ein g th a t P

32

w a s t r a n s l o c a t e d in a n o r g a n i c f o r m .

T he a ssu m p tio n

T o l b e r t a n d W ie b e

(1955) h a v e i s o l a t e d t h r e e t r a n s l o c a t i o n f o r m s o f p h o s p h a t e c o m p o u n d s ( i n ­
o r g a n i c p h o s p h a t e a n d tw o u n k n o w n p r o d u c t s ) in x y le m s a p o f b a r l e y .

M ore

r e c e n t l y M a iz e l_ e t_ a l (1956) h a s c h a r a c t e r i z e d o n e o f th e u n k n o w n s a s p h o s p f io y r l c h o l i n e .
F r o m t h e d a t a p r e s e n t e d h e r e i n , it i s n o t p o s s i b l e to a t t r i b u t e th e
d w a r f i n g e f f e c t o f th e s t o c k on t h e s c i o n t o t a l l y to d i f f e r e n c e s in t h e n u t r i ­
t i o n a l c a p a c i t i e s o f th e s t o c k .
f u lly e v a l u a t e d .

T r a n s p o r t is im p o r ta n t, but h a s not b een

T h e d i f f e r e n c e s o b s e r v e d in t h e s e tw o p r o c e s s e s m a y o n ly

b e m a n i f e s t a t i o n s o f s o m e o t h e r f a c t o r (s) m o r e d i r e c t l y c o n c e r n e d b u t
u n k n o w n to th e i n v e s t i g a t o r .

A b e t t e r u n d e rs ta n d in g of th e m e ta b o lic p r o ­

c e s s e s o f th e s t o c k a n d s c i o n a r e r e q u i r e d to m o r e f u lly u n d e r s t a n d a n d
e v a lu a te th e s to c k - s c io n i n t e r r e la tio n s h ip s .

T h e s e a im s m a y be a c h ie v e d

b y a n e v a l u a t i o n of th e m e t a b o l i c a c t i v i t i e s ( e n z y m e a n d r e s p i r a t i o n ) of th e
s t o c k a n d s c i o n a n d a u x in r e l a t i o n s h i p s .

A c r i t i c a l s tu d y o f th e r a t e o f

n u t r i e n t t r a n s p o r t , e s p e c i a l l y P 3 2 , a c r o s s g r a f t u n io n s w o u ld a l s o be in fo rm ativ e.

100

VI. SU M M A R Y

T h e e f f e c t o f t h e g r a f t u n io n a n d s t o c k - s c i o n i n t e r r e l a t i o n s h i p s , in
c e r t a i n h o r t i c u l t u r a l p l a n t s , on f o l i a r a n d r o o t a b s o r p t i o n a n d d i s t r i b u t i o n of
n u t r i e n t s w a s s t u d i e d w ith th e a i d of r a d i o a c t i v e i s o t o p e s a n d a u t o r a d i o g r a p h y .
T h e q u a n t i t y of r o o t a b s o r b e d r a d i o p h o s p h o r u s ( c p m / g m d r y w t / m g
r o o t) r e c o v e r e d in th e to p s of E a s t M a i l i n g (EM) IX, VII a n d XVI r o o t e d c u t ­
t i n g s c l o s e l y f o llo w e d s c io n v i g o r on t h e s e s t o c k s o b s e r v e d in th e fie ld .
T o p s o f E M XVI c o n t a i n e d tw i c e a s m u c h P

a s EM IX, E M V II w a s i n t e r ­

m e d ia te .
I r r e s p e c t i v e of f o l i a r o r r o o t a p p l i c a t i o n o f P 32 th e m o s t d w a r f in g
r o o t s t o c k (E M IX) a b s o r b e d t h e l e a s t , E M VII i n t e r m e d i a t e , a n d EM XVI
o r D e l i c i o u s s e e d l i n g th e l a r g e s t q u a n tity of P

32

In no c a s e w e r e s i g n i f i c a n t d i f f e r e n c e s of P

32

o r Ca

45

co n ce n tra tio n

o b s e r v e d in th e b u d u n io n s f o r m e d by M c I n to s h on EM VII, IX a n d XVI a n d
D e licio u s se e d lin g ro o ts to c k s.
No d i f f e r e n c e s w e r e o b s e r v e d b e tw e e n g e n e t i c a l l y d w a r f e d (M ac
A r t h u r 741) a n d n o n - d w a r f ( S to k e s d a le ) t o m a to p l a n t s in t h e i r a b ility to a b s o r b
fo lia r a p p lie d P

32

.

S ig n i f ic a n tl y l e s s P

32

w a s t r a n s p o r t e d to th e r o o t s of th e

d w a r f t h a n th e n o n - d w a r f p la n t.
T o m a t o p l a n t s c h e m i c a l l y d w a r f e d w ith m a l e i c h y d r a z id e a b s o r b e d
l e s s f o l i a r a p p l i e d P 3 2 , b u t no d i f f e r e n c e s in t r a n s p o r t w e r e o b s e r v e d w ith in

101

24 h o u r s a f t e r t r e a t m e n t w ith P ^ .
T h e r e w e r e no s i g n i f i c a n t d i f f e r e n c e s in th e s o i l a p p l i e d P 32 r e c o v e r e d
in th e t o p s o f g r a f t e d o r n o n - g r a f t e d t o m a t o p l a n t s .
no i n d i c a t i o n t h a t P

32

F u rth e rm o re , th e re w as

a c c u m u l a t e d a t th e g r a f t union, o r t h a t t h e u n io n i t s e l f

i n t e r f e r e d w ith th e u p w a r d t r a n s p o r t of p h o s p h o r u s .

L i k e w is e , no m a r k e d

n
,
22
35
36
45
52-54
5 5 -5 9
a c c u m u l a t i o n o f r o o t a b s o r b e d N a , S , C l , C a , Mn
, Fe
„ 64 r—
7 65 r j . 8 6
89 _ 99
140
Cu , Z n , R b , S r , M o , a n d Ba
w a s o b s e r v e d a t o r b e lo w th e
g r a f t u n io n in t o m a t o p l a n t s .
In a p p l e , C l a r k ' s d w a r f i n t e r s t e m p i e c e s o f f e r e d no r e s i s t a n c e to
th e u p w a r d t r a n s p o r t of C a

45

a p p l i e d to th e r o o t m e d i a w ith in 24 h o u r s a f t e r

tre a tm e n t.
W h e n tw o o n e - c e n t i m e t e r p h lo e m s e g m e n t s w e r e i n v e r t e d on o n e y e a r - o l d M a i l i n g I a p p le t r e e s , f o l i a r a p p l i e d P

32

a c c u m u l a t e d a b o v e th e

p h l o e m i n v e r s i o n s , w ith l e s s e r a m o u n t s a b o v e th e n o n - i n v e r t e d p h lo e m r i n g s .
S w e l l i n g s a b o v e th e i n v e r t e d p h l o e m r i n g s a n d r e d u c e d g r o w th of th e to p s
w e r e e v id en t.

A p p r o x i m a t e l y 60 p e r c e n t m o r e a d v e n t i t i o u s b u d s b r o k e on

t r e e s w ith p h l o e m i n v e r s i o n s th a n on th o s e no t h a v in g i n v e r t e d p h lo e m r i n g s .
G r a f t u n io n s f o r m e d by a p i c a l l y g r a f t i n g t o g e t h e r f o r c e d l a t e r a l s
o f v i g o r o u s l y g r o w i n g t o m a t o p l a n t s s i g n i f i c a n t l y r e d u c e d th e t r a n s p o r t of
P

32

a n d a l m o s t c o m p l e t e l y b l o c k e d th e t r a n s p o r t o f C a

45

W ith t o m a t o , s c i o n s g r a f t e d in a n i n v e r t e d p o s i t i o n g r e w l e s s in

102

t e r m s of d r y w e ig h t a n d t e r m i n a l g r o w t h .

A ls o , l e s s r a d i o c a l c i u m w a s

re­

c o v e r e d in th e i n v e r t e d s c i o n s th a n on n o r m a l l y g r a f t e d s c i o n s o r on a c o m ­
p a r a b l e s e g m e n t on a n o n - g r a f t e d p l a n t .

H ow ever, w hen e x p r e s s e d on a d ry

w e ig h t b a s i s t h e r e w e r e no s i g n i f i c a n t d i f f e r e n c e s in t h e a m o u n t s o f r o o t a b so rb ed Ca

45

r e c o v e r e d in th e s c i o n s .

B ean a n d r a d i s h l e a v e s t r e a t e d w ith C a

45

an d su b se q u e n tly a n e s th e tiz e d

w ith d i - e t h y l e t h e r t r a n s p o r t e d c o n s i d e r a b l e q u a n t i t i e s o f C a

45

f r o m th e s i t e

of a p p l i c a t i o n to a l l o t h e r p a r t s of th e p l a n t .
C h e l a t i o n of C a

45

by d i-s o d iu m e th y le n e d ia m in e te tr a a c e ta te , p ic -

r o l i n i c a c i d , 2, 5 - d i c h l o r o - 3 ,

6

-d ih y d ro x y -p -q u in o n e , and 1- a m in o - 2 -n a p th o l-

4 - s u lf o n ic a c i d d id n o t f a v o r th e b a s i p e t a l t r a n s p o r t o f C a

45

.

T he c h e la te d

c a l c i u m w a s a b s o r b e d b y th e le a f , b u t t r a n s p o r t e d o n ly t o w a r d th e p e r i p h e r y .

103

VII. GLOSSARY
A c ro p etal

In th e d ir e c tio n fro m th e tr a n s it io n zone to w a rd th e s te m
o r r o o t a p ex .

B asip etal

In th e d ir e c tio n fro m th e sh o o t o r r o o t a p e x to w a r d th e
t r a n s i t i o n zo n e.

C a rrie r free

A r a d i o i s o t o p e i s s a i d to b e c a r r i e r f r e e if i t c o n t a i n s o n ly
r a d io a c tiv e a to m s of th e e le m e n t. H o w ev er, th is id e a l is
u s u a l l y o n ly a p p r o a c h e d a n d " c a r r i e r f r e e " i s u s e d in t h i s
t e x t to m e a n no c a r r i e r a d d e d .

C lo n e

A g ro u p of in d iv id u a ls p ro d u c e d fro m a sin g le o rig in a l in ­
d i v i d u a l by a s e x u a l m e a n s .

In arch

I n a r c h i n g i s a p r o c e s s o f g r a f t i n g in w h ic h t h e s c i o n u n i t e s
w i t h th e s t o c k w h ile it is s t i l l a t t a c h e d to th e p a r e n t p l a n t .

In te rste m

A s t e m w h ic h i s u s e d to s e p a r a t e a n d c o n n e c t a r o o t s t o c k
a n d a b u d d e d o r g r a f t e d top , y e t d i f f e r e n t f r o m b o th .

L . S. D.

L e a s t sig n ific a n t d iffe re n c e .

M ic ro c u rie
(uc)

A u n it o f m e a s u r e m e n t o f r a d i o a c t i v i t y e q u a l i n g 3. 7 x 10
d isin te g ra tio n s p e r second.

R o o t-p ie c e

A ro o t w hen u s e d a s a sto ck .
to a r o o t .

R o o tsto ck

A p l a n t u s e d a s th e r o o t f o r a g r a f t e d o r b u d d e d p l a n t .

S c io n

A p l a n t u s e d a s t h e to p f o r a g r a f t e d o r b u d d e d p l a n t .

S c io n e f f e c t

A m o d i f i c a t i o n o f th e r o o t s t o c k c h a r a c t e r i n d u c e d b y th e s c i o n .

S te m -p ie c e

See i n t e r s t e m ab o v e.

S tio n

T e r m u s e d to d e s i g n a t e th e n e w i n d iv id u a l f o r m e d by g r a f t i n g
o r b u d d in g a s c i o n to th e s t o c k .

S to c k e f f e c t

A m o d i f i c a t i o n o f th e s c i o n c h a r a c t e r in d u c e d by th e r o o t s t o c k .

T B u d d in g

M e t h o d o f b u d d in g d e r i v i n g i t s n a m e b e c a u s e th e tw o c u t s m a d e
on t h e s t o c k i n t e r s e c t so a s to f o r m a " T " .

4

T h e scio n is g ra fte d d ire c tly

104

V III. L I T E R A T U R E C IT E D

1

. A s s o c i a t i o n o f O f f ic ia l A g r i c u l t u r a l C h e m i s t .
A n a l y s i s . S e v e n th E d itio n .

1950.

O ffic ia l M e th o d s of

2. A m o s , J. , T . N. H o bly n, R. J. G a r n e r a n d A. W. W itt. 1936. S tu d ie s
in i n c o m p a t i b i l i t y o f s t o c k a n d s c io n . I. I n f o r m a tio n a c c u m u l a t e d d u r in g
t w e n ty y e a r s of t e s t i n g f r u i t t r e e r o o t s t o c k s w ith v a r i o u s s c io n v a r i e t i e s
a t E a s t M a il in g . E a s t M a i l i n g R e s . S ta. Ann. R e p t. fo r 1935: 8 1 -8 9 .
3. A r g l e s , G. K. 1937. A r e v i e w o f th e l i t e r a t u r e on s t o c k - s c i o n i n c o m p a b i l i t y in f r u i t t r e e s , w ith p a r t i c u l a r r e f e r e n c e to p o m e a n d s to n e f r u i t s .
T e c h . C o m m u n i c a t i o n , No. 9 Im p. Bur. F r u i t P r o d . E a s t M a ilin g .
4. A r n o n , D. I . , P. R. S to u t a n d F . S ip o s . 1940. R a d i o a c tiv e p h o s p h o r u s
a s an i n d i c a t o r o f p h o s p h o r u s a b s o r p t i o n of to m a to f r u i t s a t v a r i o u s
s t a g e s of d e v e l o p m e n t . A m e r . j o u r . Bot. 27: 7 9 1 -7 9 8 .
5. A s e n , S . , S. H. W i t t w e r a n d O. N. H i n s v a r k . 1953. F o l i a r 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 r a d i o p h o s p h o r u s by C h r y s a n t h e m u m m o r i f o liu m .
P r o c . A m e r . S oc. H o r t . S ci. 62: 4 6 6 -4 7 0 .
6

. B a ile y , L. H. 1949. M a n u a l o f c u l t i v a t e d p l a n t s .
M a c m i l l a n , N ew Y o r k .

R e v i s e d e d . , p. 515.

7. B e a k b a n e , A. B. , a n d M. E . R e n w ic k . 1936. A p r e l i m i n a r y r e p o r t on
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