A study of the comparative value of Juniperus scopulorum and Juniperus virginiana as understock for
four juniper clones
by Neil J Van Sloun
A THESIS Submitted to the Graduate Faculty in partial fulfillment of the requirements for the degree
of Master of Science in Horticulture at Montana State College
Montana State University
© Copyright by Neil J Van Sloun (1959)
Abstract:
No significant difference appears to exist between Juniperus sobpu-lorum and Juniperus Virginiana as
understocks for J. chinesis pfitzeriana, J. hOrizqntalis lividus, J. sabina tamariscifolia, or J. scopulorum
Mon-tana. No, 1. Successful unions were established for all grafts in which the bark of the scion and
stock was close enough to permit the establishment of a callus bridge. Two somewhat different
developmental sequences occur, depending upon the distance separating scion from stock, Tfflien scion
and stock are relatively near one another' the sequence involves (l) the formation of a callus bridge
followed by (2) the development of a cambial bridge, and (3) the subsequent production of new
secondary xylem and secondary phloem. When scion and stock are relatively far apart a union of the
vascular tissues of scion and stock precedes cambial activity through the differentiation of some callus
tissue into tracheids and sieve cells on either side of the newly formed cambium. Parenchyma cells of
the cortex, phloem, and xylem, including the phloem and xylem rays, contributed to callus formation;
the cambium contributed little, if at all. A STUDY OF
THE COMPARATIVE VALUE'OF JUHIPERUS
SCOPULORUM AHD JUHIPERUS VIRGIHIAHA AS UNDER­
STOCK FOR FOUR JUNIPER CLONES
ty
HEIL J. VAH SLOUH
A THESIS
Subm itted t o t h e Graduate FAcuIty
in
p a r t i a l f u lf il lm e n t o f th e requirem ents
f o r th e d eg ree o f
M aster o f S c ie n c e in H o r tic u ltu r e
at
Montana S t a t e C o lleg e
Approved:
______________
Head, Major Department
Chairman, Exam ining Committee
B'o;zeman:.>J(i.Hont:Ana;i ■ .
Decemher;, 1959
N 51?
V 314 s
-
2-
ACKNOWLEDGEMENTS
The au th or w ish es to extend h is s in c e r e a p p r e c ia tio n t o Dr. V incent
I v e r s on. Dr. Ray E v ert, and t o Mr. Leonard Yager f o r t h e i r v a lu a b le
a s s is t a n c e and s u g g e s tio n s throughout th e co u rse o f t h i s stu d y .
139171
-3 “
TABLE OF COETEWTS
Page
ACEEOWLEDOEMEET ...................................................................................................................
”2
TABLE OF COETEETS ..............................................
3 .
ABSTRACT............................................................
Li.
IET RODEC T IO E .......................................... .... ........................................................... . . .
5
LITERATURE REVIEW...............................................................................................................
5
MATERIALS AED" METHODS......................................................................................................
8
RESULTS . . . . . . .
.................................................................................................
.
11
DISCUSSIOE 'AED COECLUSIOES..............................................................................................l6
LITERATURE C I T E D ................................................................
PLATES
..................................................
22
25
ABSTRACT
Ho s i g n i f i c a n t d if f e r e n c e appears t o e x i s t betw een J u n ip e r u s"sobpulorum and Juniperus V ir g in ia n a as u n d ersto ck s ' f o r J e ch in es i s p 'fitz eria'na,
J 6 h O r iz q n ta lis I i v id u s , J 6 sa b in a t a m a r is c i f d l ia , or J6 scopulorum Mon-""
ta n a '. Ho6 I 6 S u c c e s s f u l u n ion s were e s t a b lis h e d f o r a l T "grafts in w hich
th e bark o f th e s c io n and s t o c k was c l o s e enough t o "permit th e e s t a b lis h ­
ment ^o f a c a l l u s b r id g e . Two somewhat d if f e r e n t d evelopm en tal sequences
occu r, depending upon th e d is ta n c e s e p a r a tin g s c io n from s t o c k , Tfflien
s c io n and s to c k a r e r e l a t i v e l y n ea r one a n o th e r 'th e sequence in v o lv e s
( l ) th e fo rm a tio n o f a c a llu s b rid g e fo llo w e d by (2 ) th e developm ent o f
a cam bial b r id g e , and ( 3 ) th e su bseq u en t p ro d u ctio n o f new secondary
xylem and seco n d a ry phloem* Tfflien s c io n and s t o c k a re r e l a t i v e l y fa r
a p art a un ion o f t h e v a s c u la r t i s s u e s o f s c io n and s to c k p reced es cam bial
a c t i v i t y th rou gh th e d i f f e r e n t i a t i o n o f some c a llu s t i s s u e in t o tr a c h e id s
and s ie v e c e l l s on e i t h e r s id e o f th e new ly formed cambium. Parenchyma
c e l l s o f th e c o r t e x , phloem, and xylem , in c lu d in g th e phloem and xylem
r a y s , c o n tr ib u te d t o c a llu s fo m a tio n ? th e cambium c o n tr ib u te d l i t t l e ,
i f at a ll .
INTRODUCTION
Ju n ip ers are propagated Uy s e e d s , la y e r s , c u t t in g s , and g r a f t s „
S p e c ie s th a t ro o t w ith d i f f i c u l t y or do n o t grow tr u e t o p a ren t ty p e from
seed are propagated by g r a f t in g .
p la n t i s d e s ir e d more q u ic k ly ,
G ra ftin g is a ls o used when a s a la b le
At p r e s e n t, two u n d erstock s a re commonly
used i n ju n ip e r g r a f t in g , Juniperus vj.rgihiq.na- and Juniperus c h in e n s is .
Juniperus soopulorum i s n a t iv e to th e "Rocky Mountain area and o ffe r s more
h a rd in ess and drought r e s is t a n c e than _J, v ir g in ia n a or_J, c h in e n s is ,
The
aim o f t h i s in v e s t ig a t i o n was t o d eterm ine w hether J* scopulorum compares
fa v o r a b ly w ith J , v ir g in ia n a as an u n d ersto ck in th e com m ercial propaga­
t io n o f ju n ip e r s .
LITERATURE REVIEW
G ra ftin g o f p lan s i s a v ery a n c ie n t a r t , d a tin g back t o 1560 B, C,
(3 b )» There are many methods o f g r a fta g e ( I , 17, 19, 27} and•many u ses
and reason s f o r g r a f t in g (3 b )«
G rafts have been s u c c e s s f u l on a lg a e ,
m o sses, low er v a s c u la r p la n ts ( S e l a g i n e l l a ) , herbaceous monocots and
d i c o t s , and woody gymnosp'erms- and angiosperm s ( 3b) «
Many i n t e r - s p e c i f i c
and in t e r - g e n e r ic g r a fts have been s u c c e s s f u l b u t s u c c e s s f u l g r a fts betw een
f a m ilie s are q u ite rare ( 3b) »
G r a ftin g o f ju n ip e r s has been done f o r many y e a r s ,
M allin son (2 8 ) in
1926 d e sc r ib e d th e g r a f t in g procedure f o r ju n ip ers and m entioned J. v i r g i ­
n ian a as th e u n d e r sto c k .
S in ce then J , o h in en si s , J , e z o e ls a s t r i c t a , _J»
communis h ib e m ic a , J , h o r iz o n t a lis plumosa , and Thuja o r i e n t a l i s have b een
n oted in th e l i t e r a t u r e ( 5 , 6 , 7 , 15, 2 l) as u n derstocks in ju n ip er propaga­
tio n ,
At p r e s e n t J , v ir g in ia n a and
c h in e n s is are most h ig h ly recommended
( 19, 21, 22, 27, 39, Uo) as u n d e r sto c k s.
-
6-
A lthough g r a f t in g i s a v ery a n c ie n t a r t , a n atom ical s t u d ie s o f g r a f t
union developm ent d a te back m erely t o th e m id -e ig h te e n th cen tu ry ( 1 2 ) .
S in ce th e n , i n t e r e s t in g r a f t in g has proceeded a lo n g two l i n e s ;
(a )
a p p lie d u s a g e s - in a g r ic u lt u r e and (b ) a n a to m ica l and p h y s io lo g ic a l s t u d i e s ,
w ith n e it h e r area c o n tr ib u tin g s i g n i f i c a n t l y to an u n d ersta n d in g o f a l l
fa c to r s in v o lv e d in th e un ion o f s t o c k and scio n #
The c h r o n o lo g ic a l seq u en ce o f o n t o g e n e t ic . phases in th e e sta b lish m en t
o f th e g r a ft u n ion has b een summarized by Hartmann and K ester (19) a s ;
"(a) e s ta b lish m e n t o f in tim a te co n ta c t o f a co n sid e r a b le amount o f th e
cam bial reg io n o f b oth s to c k and s c io n under fa v o r a b le en vironm en tal con­
d i t i o n s , (b) p ro d u ctio n and in t e r lo c k in g o f parenchyma c e l l s ( c a llu s t i s s u e )
o f b oth s to c k and s c io n , (c ) p ro d u ctio n o f new cambium a c r o ss th e c a llu s
'b r id g e 1', and (d) form ation o f new xylem and phloem from th e new v a s c u la r
cambium in th e c a llu s b r id g e " .^
A s l i g h t l y d if f e r e n t developm en tal seq uence has been rep o rted as
o ccu rr in g in F ic o tia n a ( 1 0 ) .
Here s ie v e elem ents and xylem elem ents d i f ­
f e r e n t i a t e w ith in th e c a llu s b rid g e co n co m ita n tly w ith th e developm ent o f
cam bial t i s s u e , thus g iv in g r i s e to a union o f th e younger v a sc u la r t i s s u e s
o f s c io n and s t o c k p r io r t o cam bial a c t i v i t y .
Sass (35) su g g este d t h a t a
s im ila r p r o c e ss occurs in ap p le when poor m atching r e s u lt s in e x c e s s iv e
c a llu s fo rm a tio n .
Frequent sta tem en ts are made in t h e o r e t i c a l and p r a c t ic a l g r a ftin g
li t e r a t u r e th a t o n ly th e cambium p o s s e s s e s p r o l i f e r a t i v e c a p a c i t i e s .
^Hartmann, H. T„, and D. E. K e ster .
p le s and p r a c t i c e s . P r e n t ic e - H a ll, Inc#
275»
TL
Fames
1959» P la n t "propagation, p r in c i­
Englewood C l i f f s , F, J. Pp. 273-
-
and MaoDaniels (1 3 ) s t a t e
7"
„ e budding and g r a f t in g have as t h e i r b a s is
th e a b i l i t y of th e cambium o f b o th s t o c k and s c io n to d ev elo p c a l l u s , ,
This i s co n tra ry to th e ev id e n c e forwarded by a m a jo r ity o f in v e s t ig a t o r s
( 3 , b, 2i|, 3 1 » 35» 36) in d ic a t in g t h a t th e cambium
o f t e n p la y s a su b o rd in a te
r o le or i s n o t th e o n ly t i s s u e reg io n in v o lv e d in c a llu s i n i t i a t i o n .
Paren­
chyma c e l l s o f th e c o r te x ( 10, Elf., 37)» phloem (bs 2^» 26» I).I, 36, 10) phloem
rays (I+, 2l|.» 3 1 » 36)» m ed u llary rays (18» 3&)» xylem ( 10» 36)» immature
.xylem rays (l+, 3 6 » 37)» xylem rays (J4) , and p it h ( 2l+» 37» 10) , have been men­
tio n e d as c o n tr ib u to r s t o c a llu s t i s s u e ,
At l e a s t one in v e s t ig a t o r s t a t e d
th a t th e cambium c o n tr ib u te d v ery l i t t l e to c a llu s i n i t i a t i o n (3 5 )»
I t is
obvious th a t c o n tro v er sy e x i s t s With regard t o th e r e l a t i v e r o le s played by
th e cambium and o th e r t i s s u e s in c a llu s fo n r a t io n ,
Some d isa g reem en t appears to e x i s t a l s o , as to th e r e l a t i v e r o le s
p layed by th e s c io n and sto d k in th e form ation o f c a llu s t i s s u e ,
K o sto ff
( 23) rep orted th a t in th e S o la n a cea e c a llu s t i s s u e j o in in g th e s c io n and
s to c k i s c h i e f l y th e product o f t h e s t o c k .
This sta tem en t i s n ot in a g ree­
ment w ith o b se r v a tio n s on o th e r p la n ts rep o rted by th e m a jo r ity o f w ork ers,
who s ta te d t h a t c o n tr ib u tio n s from s t o c k and th e s c io n a r e app roxim ately
e q u a l.
In a d d it io n to a c o n s id e r a tio n o f th e sequence o f e v e n ts in th e form a­
t i o n o f s u c c e s s f u l g r a ft u n io n s» one must attem pt t o determ ine why c e r t a in
u n ion s f a i l .
S e v e r a l f a c t o r s have been su g g ested in th e l i t e r a t u r e as
b e in g r e s p o n s ib le f o r g r a f t union f a i l u r e s .
They a r e s
wood d is c o n t in u it y
2Eames, A. J,» and L, H„ M acD aniels, 19U7» In tr o d u c tio n t o p la n t
anatomy, McGraw-Hill Book C o,, I n c , , Hew ^ork6 Pp, 20 1 ,
-
8 «-
( 2, 8 S 32, 3 8 )» xylem d is t o r t io n ( 33) , e x c e s s iv e c a llu s form ation ( 8 , 3 2 ,
33) , s e a s o n a l r e g r a f ts r e s u lt in g in e x c e s s iv e c a llu s form ation ( o fte n
su b e r iz e d ) ( 3) ,
cambium d e s tr u c tio n ( 20) , bark in t e r r u p t io n ( 20) , phloem
d is c o n t in u it y ( 3) , and uneven s ta r c h b a la n c e ( 8 , 20, 25, 30, 33)»
MATERIALS AHD METHODS
This in v e s t ig a t i o n was conducted a t Montana S ta te C o lleg e from Novem­
b e r 1958 t o November 1959®
The p la n t m a te r ia ls u sed were _Je scopulorum ( Common 1R ocky Mountain
ju n ip e r ) and J e v ir g in ia n a (E a stern Red Cedar) as u n d e r sto c k s, and Je
c h in e n s is p f it z e r ia n a ( P f i t z e r j u n ip e r ) , J e h o r iz o n t a lis I i vid u s ( L ivid us
j u n ip e f ) , Je sa b in a t a m a r is c i f o lia (S a v in ju n ip e r , "Tam" v a r ie t y ) , and _Je
scopulorum , Montana No* I (Montana Noe I ju n ip er* ) as s c Ionwoode
Juniperus scopulorum u n d ersto ck s were ob tain ed from th e F o r e st N prsery
o f Montana S ta te U n iv e r s it y a t M isso u la , Montana *
These u n d ersto ck s a v er­
aged 3 /1 6 in ch in d ia m eter a t th e b a se and f i v e in ch es in h eig h t*
Juniperus
v ir g in ia n a u n d ersto ck s were o b ta in ed from th e P lu m field N u r se ries a t Fremont,
N ebraska,
These u n d ersto ck s averaged l / k in c h in d ia m eter at th e b a se and
e ig h t in ch es - in h eig h t*
The _£* Scopulorum u n d erstock s were 2 -1 (2 y ea r o ld ,
once tr a n s p la n te d ) s e e d li n g s , th e U0 v ir g in ia n a n o n -tr a n sp la n te d 2 -y e a r o ld
s e e d lin g s *
The s c io n s were o b ta in ed from p la n ts growing on th e Montana
S ta te C o lleg e campus and in th e Bozeman a r e a .
The _J, c h in e n si s p f it z e r ia n a
s c io n s averaged 11 in ch es in le n g th , th e o th e r th r e e v a r i e t i e s s i x in ch es
* S p e c ia l c lo n a l s e c t io n h avin g u p r ig h t form and b lu e c o lo r made and
propogated by P lu m field N u r s e r ie s , Fremont, Nebraska*
in length#'
A ll th e s c io n s were o f th e cu rren t yearns grow th.
The s c io n s
were cu t two days p r io r t o g r a f t in g ( e a r ly January, 1959) and kept r e f r ig e r ­
ated f o r t h i s p e r io d under c o n d itio n s o f h ig h r e l a t iv e h u m id ity ,
' The same s o i l was u sed th rou gh out th e in v e s t ig a t i o n ,
p a rts loam to p s o i l ,
A m ixture o f 5
I p a rt sand, and I p a rt manure was u s e d .
Four hundred jJ„ scopulorum s e e d lin g s were p o tted on 21 November, 1958
in Ip-inch p l a s t i c p o t s ,
The p la n ts were sto r e d in a c o o l greenhouse (m in i­
mum tem perature o f 5 5° F) u n t i l th e y were g ra fted in January 1959«
hundred and f i f t y
Four
v ir g in ia n a s e e d lin g s were p o tte d on 18 December, 1958
in !+-inch p l a s t i c p o ts and handled as th e o th er s e e d li n g s .
J0 v ir g in ia n a s e e d lin g s were trimmed b e fo r e p o t t in g .
Roots o f th e
Those o f th e _J,
scopulorum s e e d lin g s were n o t trimmed b eca u se t h e i r ro o t system s were n ot
s o la r g e as t h o s e o f th e _J, v ir g in ia n a s e e d lin g s .
On 10 January, 1959 th e u n d ersto ck s were so rted t o a ch ie v e an "even"
u n derstock*
One hundred and f o r t y J» scopulorum and f i f t y J , v ir g in ia n a
s e e d lin g s were d isca rd ed b ecau se o f low v i t a l i t y and crooked stem s.
The s id e g r a f t was u sed throughout t h i s in v e s t ig a t i o n .
An o b liq u e
i n c i s i o n Ig- in c h e s lon g and p e n e tr a tin g 1 /3 o f th e d iam eter o f th e stem
was made on th e low er two in ch es o f th e u n d e rsto c k .
I n to t h i s in c i s i o n was
in s e r t e d th e low er end o f th e s c io n t h a t had been cu t to form a wedge Igin c h e s lo n g .
The s c io n was p laced as f a r as p o s s ib le in t o th e u n d ersto ck .
Care was ta k en t o li n e up as c l o s e l y as p o s s ib le th e cam bial reg io n s o f
s c io n and s t o c k .
s tr ip s .
The g r a f t was th en bound s e c u r e ly w ith rubber budding
The tim e o f g r a f t in g and th e numbers o f each g r a ft com bination
a re l i s t e d in Table I ,
-
10 -
Table I - G raft com binations and tim e o f g r a f t in g .
Graft com binations
S to ck
S cion
J , v ir g in ia n a
P fitz e r
J , scopulorum
P fitz e r
J , v ir g in ia n a
L ivid us
L ivid u s
J , scopulorum
J , V ir g in ia n s
Montana No, I
J, scopulorum
Montana No, I
J , V ir g in ia n s
S avin
J , scopulorum
S avin
Number
made
100
'80
100
60
100
60
100
60
12
15"
16
16
18
18
23
2l|.
Date
made
Jan 59
Jan 59
Jan 59
Jah 59
Jah 59
Jan 59
Jan 59
Jan 59
On th e same day th e g r a fts were made, th e p la n ts were p la ced under
in t e r m it t e n t m ist in a greenhouse a t a minimum tem perature o f 6 5 ° F6
The
au tom atic m is t system was o f an llIn=Bed" ty p e w ith Type A-6 "Humid o-m ist"
n o z z le s ( s e l f - c l e a n i n g ) .
The m ist was c o n tr o lle d by an " e le c tr o n ic le a f " .
The J , c h in e n s is p f it z e r ia n a g r a fts were kept under' th e m ist a t 65° F 0 fo r
te n d ays, th e o th e r g r a f t s f o r fo u r te e n d a y s.
A fte r t h i s treatm en t th e
g r a fts were p la ced in a d if f e r e n t greenhouse a t 55° F,
At t h i s tim e m o ist
sphagnum moss was packed around each g r a ft t o preven t th e g r a fts from
d ryin g o u t.
On 26 February, ^O.: J,. h o r iz 'o n ta lis llv id u s g r a fts on 1J,
■
.
v ir g in !a n a were moved back t o t h e 6 5 °" F, house and fo r c e d .
A lso moved
back t o th e 65° F , house w e r e _ fiv e each o f th e o th er g r a ft co m b in a tio n s.
A ll o f th e above s c io n s were a l i v e and q u ite green .
On 15 February, on e-
h a lf o f th e to p o f th e u n d e rsto c k was removed on a l l th e p la n t s .
rem aining h a l f was removed on 25 March,
The
On 2? J u ly , th e growing g r a fts
were counted and t h e p la n ts s e t out in n u rsery fram es.
An a n a ly s is o f
v a r ia n c e t e s t was conducted to determ ine any s i g n i f i c a n t d if f e r e n c e s b e­
tw een th e two u n d e r sto c k s.
- I l-
The c o l l e c t i o n o f m a te r ia l f o r an a n a to m ica l stu d y o f th e g r a ft u n ion s
was begun on 26 February 1959,
At t h i s tim e , one g r a ft o f each s t o c k - s c io n
com bination was removed and p la ced in Craf I I I s o lu t io n ( 2 3 ) e
jJ, h o r iz o n t a l! s I i v id u s on
(2 3 )»
An a d d it io n a l
v i r g in ia n a g r a f t was p la ced in FAA s o lu t io n
T h erea fte r two J e h o r iz o n t a lis liv id u s on _Je v ir g in ia n a g r a fts were
f ix e d ( one in Craf I I I and one in FAA) a't w eek ly in t e r v a l s u n t i l 21 May,
One" each o f th e sev en o th e r com binations were fix e d in Craf I I I a t m onthly
in te r v a ls .
A fte r t h i s tr e a tm e n t, a l l sam ples were s o fte n e d in a m ixture
1
o f one p a rt fuming h y d r o flu o r ic a c id and one p art
fo r one w eek.
p e r c e n t e t h y l a lc o h o l
They w ere th e n dehydrated and embedded i n c e l l o i d i n ( 2 3 ) ,
L o n g itu d in a l and tr a n s v e r s e s e c t io n s were cu t a t 15 m icrons w ith
a s l i d i n g m icrotom e.
They were th en s ta in e d w ith ani I in b lu e and io d in e -
p o ta ss iu m -io d id e and mounted in g ly c e r in e ,
EESULTS
G ra ftin g S u c c e s s ,
The a n a ly s is o f v a ria n c e ( Table IIB ) in d ic a t e s t h a t no s i g n i f i c a n t
d if f e r e n c e e x is t e d betw een t h e two u n d e r sto c k s, J , soopulorum and J , v i r g i ­
n ia n a ,
B ecause o f th e slo w growth o f j u n ip e r s , t o t a l lin e a r growth o f th e
d if f e r e n t s t o c k - s c io n com binations was n o t m easured.
The g r a fte d p la n ts
were sep a ra ted in t o two groups $ dead and s u r v iv in g ( s u r v iv in g b ein g d e fin e d
as th o s e p la n ts show ing v i s i b l e s ig n s o f grow th ),
A ta b u la t io n o f s u r v iv a l p ercen ta g es i s p resen ted in Table IIA ,
The
S avin on J , soopulorum e x h ib ite d 68 p erce n t s u r v iv a l w h ile Savin on J ,
v i r g in ia n a r e s u lt e d in Lf.3 p ercen t s u r v iv a l,
L ivid us on J , v i rg in ia n a and
J, soopulorum was ap p roxim ately eq u al w ith 52 and J4.3 p e r c e n t, r e s p e c t i v e ly .
-
12 -
Table IIA - P ercen ta g e o f grow ing s t o c k - s c io n com binations on
. .. . .
27 J u ly ,. .195%...................................... : .................................
Graft com binations
S tock
S cio n
L ivid us
J . scopulorum
L ivid u s
J . virginiana"
S avin
J . scopulorum'
S avin
J . v ir g in ia n a
P fitz e r
J e scopulorum
P fitz e r
J . v ir g in ia n a
Montana Ho. I
J . scopulorum
Montana h o . I
J e v ir g in ia n a
P ercen t
S u rv iv in g
k3
■52
68
h3
0
I
21
15
T able IIB - A n a ly s is o f v a ria n c e f o r d ata in Table IIA .
S .S .
D .F.
Treatment
Error
I
6
T otal
7
'
M.S.
7 5 .5 3
7 5 .3 5
I1.259. l 6
7 0 9 .8 6
C alc.
.1 0 6
F. *
Tab, '
5 .9 9
1 5 .7 5
Ii332i.li9
*F v a lu e i s n o t s i g n i f i c a n t .
There was l i t t l e d if f e r e n c e in growth among th e s e v e r a l g r a ft combina­
t io n s ( P la t e l ) .
I t was n oted t h a t some g r a f t s w hich appeared t o have
h ea le d and formed c a llu s t i s s u e remained green and a l i v e fo r some tim e b u t
e v e n tu a lly d ie d .
Montana No. I u n ite d alm ost tw ic e as w e ll w ith J . scopulorum than w ith
J . V ir g in ia n a y a lth o u g h b o th p erce n ta g e s were q u ite low ( Table I IA )«
d id v ery p o o r ly on b o th u n d e r sto c k s„
P fitz e r
-
13 -
G eneral D e s c r ip tio n o f th e Stem,
The com p lete 'stem (b o th J6 scopulorum and _J, v ir g in ia n a ) c o n s is t s o f
periderm , c o r te x , o b lit e r a t e d prim ary phloem, s e v e r a l increm ents o f se c o n ­
dary phloem , th e oam bial r e g io n , s e v e r a l increm ents o f seco n d a ry xylem
and p i t h .
The c o r te x i s composed e n t i r e l y o f s t a r c h - a n d /or r e s in - c o n t a in in g
parenchyma c e l l s and p o s s e s s e s numerous in t e r c e l l u l a r s p a c e s .
The elem en ts in th e v e r t i c a l system o f th e secondary phloem are
arranged t a n g e n t ia ll y and a lt e r n a t e w ith one an oth er in th e seq u en ceJ
s ie v e c e l l s , parenchyma c e l l s , s ie v e c e l l s , parenchyma c e l l s , parenchyma
c e l l s , f i b e r s , e t c , ( P la t e I I , A ),
The parenchyma c e l l s o f th e parenchyma
stra n d s and o f th e rays c o n ta in s ta r c h a n d /o r “r e s in o u s ” m a te r ia l ( 9 )»
T a n g e n tia lly e lo n g a ted r e sin , ca n a ls surrounded by e p i t h e l i a l c e l l s are
found i n th e n o n -fu n c tio n a l phloem .
The v e r t i c a l system o f th e secon d ary xylem c o n s is t s o f tr a c h e id s and
gu m -con tain in g parenchyma c e l l s .
The xylem rays ( h o r iz o n t a l system ) are
made up e x c l u s i v e ly o f s t a r c h - an d /o r r e s in - c o n t a in in g parenchyma c e l l s .
A ll parenchyma c e l l s ( o f c o r te x , p h lo e m ,'xylem, and p ith ) p o sse ss
prim ary w a lls .
The S u c c e s s f u l G r a fts,
T w enty-four o f fo r ty -tw o g r a fts stu d ie d a n a to m ic a lly e x h ib ite d su c ­
c e s s f u l u n io n s ,
In each o f th e tw e n ty -fo u r s u c c e s s f u l g r a f t s th e s c io n
had been in s e r t e d in t o th e s to c k in such a manner t h a t th e bark o f th e
s c io n e i t h e r came in to c o n ta c t w ith t h a t o f th e s to c k or n e a r ly so*
A ll
parenchyma c e l l s o f th e stem seem t o be e q u a lly cap ab le o f p r o li f e r a t io n
and form ation o f c a llu s t i s s u e , w ith th e p o s s ib le e x c e p tio n o f th o se in
th e p ith *
Eo n o ta b le d if f e r e n c e s appear t o e x i s t in th e b eh a v io r o f th e
v a rio u s com binations o f s c id n and s t o c k , nor in t h e i r s ta r c h c o n te n t.
The seq u en ce o f ev en ts in th e un ion o f s c io n and s t o c k was s im p le s t
in th o se g r a f t s w ith a more a ccu ra te align m en t o f t i s s u e s o f s c io n and
s to c k , th a t i s , w ith o u te r bark a d ja c e n t t o o u te r bark, in n e r bark a d ja ­
cen t t o in n e r bark, e t c , ( P la t e I I I , A ),
The o n to g e n e tic sequence in c lu d e s
( l ) th e p r o li f e r a t io n o f parenchyma o f th e c o r te x (b o th phloem and ray p ar­
enchyma) o f th e bark fo llo w e d by (2 ) th e developm ent o f a cambium ( P la t e
III,
A, a t c) in th e new ly formed t i s s u e r e s u lt in g in th e union o f th e
cambia o f s c io n and s t o c k , and (3 ) th e su bseq u en t p ro d u ctio n o f new
secondary phloem and secon d ary xylem .
The sequence o f ev en ts in th e g r a f t s th a t p o s se s s e d a l e s s a ccu ra te .
align m en t o f t i s s u e s o f s c io n and bark ( e , g , , w ith o u te r bark o f th e
s c io n a d ja c e n t t o in n e r bark o f th e s to c k and in n er la r k o f th e s c io n
a d ja cen t t o secon d ary xylem o f th e s to c k ) i s s im ila r t o t h a t m entioned
above*
In th e second ty p e g r a f t , how ever, th e parenchyma (b o th xylem and
ray parenchyma) o f th e secon d ary xylem ( P la t e I I I , B) o f th e s to c k , in
a d d itio n t o th e parenchyma o f th e bark o f b o th s c io n and s to c k , p r o li f e r a t e
and c o n tr ib u te t o t h e form ation o f th e c a llu s t i s s u e in w h ich a cam bial
b r id g e e v e n t u a lly d if f e r e n t i a t e s *
This ty p e union o f s c io n and sto c k r e s u lt s
in th e form ation o f a U -shaped cam bial u n ion ( P la t e I I I ,
C at c )*
I f s c io n and s to c k are in a c tu a l c o n ta c t w ith one a n o th er or n e a r ly
s o , a c a m tia l union i s a ch iev ed w ith a r e l a t i v e l y s h o r t p erio d o f tim e .
-
15 -
a l l new v a s c u la r elem ents ( s i e v e c e l l s and t r a c h e id s ) r e s u lt from cam bial
'
a c t i v i t y and th u s have t h e i r lon g axes o r ie n te d v e r t i c a l l y .
I f on th e
o th e r hand s c io n and s t o c k are r e l a t i v e l y f a r ap art and thus req u ire a
r e l a t i v e l y la r g e c a llu s b r id g e , tr a c h e id s and sieve- c e l l s w ith t h e i r long
axes o r ie n te d h o r iz o n t a lly may d i f f e r e n t i a t e on e i t h e r s id e o f and p r io r
t o a c t i v i t y o f th e new ly formed cam bial b r id g e ( P la t e IV, C),
Under such
circu m stan ces a un ion o f th e xylem and phloem o f th e s c io n and t h e sto c k
p reced es cam bial a c t i v i t y w ith in th e cam bial b r id g e .
In most o f th e s u c c e s s f u l g r a f t s th e bark o f o n ly one s id e o f th e
s c io n was a lig n e d w ith th a t o f th e s to c k .
This i s due t o th e f a c t th a t
most o f th e s c io n s are s m a lle r than th e area o f th e s t o c k in which th ey
were in s e r t e d ,
As a r e s u l t , n o t o n ly i s th e r e a sp ace b etw een b oth s id e s
o f th e s c io n and th e s to c k , b u t a ls o betw een th e two h a lv e s o f th e s to c k
n o t sep a ra ted by th e s c io n ( P la t e IV, A),
I f th e d is ta n c e between s c io n
and stock" i s n ot v ery g r e a t, th e s e a rea s become occupied by c a llu s t i s s u e
th a t remains parenchym atous.
I f , on th e o th e r hand, a r e l a t i v e l y la r g e
gap e x i s t s betw een s c io n and s t o c k , th e c a llu s t i s s u e b r id g in g th e s e a reas
may g iv e r i s e t o a cambium t h a t runs p a r a l l e l t o th e s c io n and another th a t
runs p a r a l l e l to th e s to c k ( P la t e IV, B )„
The cambium ly in g a d ja cen t t o th e
s c io n e v e n tu a lly u n it e s w ith th e cambia a t e it h e r end o f th e s c io n , and th e
cambium ly in g a d ja cen t t o th e s to c k w ith th e p r e - e x i s t i n g cambium o f th e
sto c k .
I f th e d is t a n c e s are r e l a t i v e l y g rea t between b o th h a lv es o f th e
s to c k an d 'th e s c io n , a com plete r in g o f cambium may d ev elo p around th e
s c io n and around each h a l f o f th e s to c k , _ These cambia su b seq u en tly g iv e
r i s e t o secon d ary xylem and secon d ary phloem .
Very commonly groups o f
more or l e s s o b liq u e ly o r ie n te d tr a c h e id s d i f f e r e n t i a t e from th e c a llu s
t i s s u e th a t b r id g e s th e s c io n and s t o c k r e s u lt i n g in a u n io n o f th e xylem
o f th e two p a r ts o f th e g r a f t ( P la t e I I , B )„
The U n s u c c e ss fu l G r a fts»
In sev en teen o f t h e 'e ig h t e e n u n s u c c e s s f u l g r a fts stu d ie d a n a to m ic a lly ,
th e s c io n was embedded e n t ir e ly in th e secon d ary xylem o f th e s to c k ( P la t e
I V ,' D) $ in o th e r words th e r e was no a lign m en t o f th e bark o f th e s c io n w ith
th a t o f th e s t o c k .
C allus t i s s u e produced b y b o th s c io n and sto c k gave
r i s e t o e i t h e r s c a r t i s s u e or a periderm b e fo r e a b rid g e cou ld be a ch iev ed
th ereb y p r e v e n tin g th e un ion o f t h e s c io n w ith th e s t o c k .
The o th er unsuc­
c e s s f u l g r a ft examined a n a to m ic a lly ( P f i t z e r on J 0 scopulorum ) had a n e a r ly
p e r fe c t alignm ent o f th e t i s s u e s o f s c io n and s t o c k , b u t l i t t l e or no pro­
l i f e r a t i o n o f parenchyma had occurred ( s e e comments on P f i t z e r com binations
in D is c u ss io n s and C o n c lu s io n s),
DISCUSSION MD CONCLUSIONS
As in d ic a te d in th e in tr o d u c tio n , th e aim o f t h is in v e s t ig a t i o n was
t o determ ine w hether Juniperus scopulorum compares fa v o ra b ly w ith _£» V ir g i­
n ia n a as an u n d e rsto c k in th e com m ercial p rop agation o f ju n ip e r s ,
_J,
v ir g in !a n a and J , scopulorum are alm ost i d e n t i c a l a n a to m ic a lly and b o th
have th e same chromosome number, 2n=22 (1 1 , 2 9 ) ,
For t h e s e reasons one
m ight exp ect b o th s p e c ie s t o prove e q u a lly s u it a b le as u n d ersto ck s fo r
d ip lo id ju n ip e r s .
The s t a t i s t i c a l a n a ly s is o f a l l d a ta in d ic a t e s t h a t no s i g n i f i c a n t
d if f e r e n c e e x i s t s betw een th e perform ance o f J, scopulorum and J, v ir g in ia n a
-
17 -
as u n d ersto ck s f o r th e m a te r ia ls stu d ied *
O bviously con tin u ed o b se r v a tio n s
over a lo n g e r p erio d o f tim e would be d e s ir a b le t o determ ine w ith c e r t a in t y
th e t r u e v a lu e o f t h e s e two u n d erstock s*
During t h i s p e r io d such f a c t o r s
as ro o t fo rm a tio n , w in te r in ju r y , h a r d in e s s , drought r e s is t a n c e , g en era l
growth h a b it s , d is e a s e r e s is t a n c e , and p o s s ib le dw arfing e f f e c t s sh ould be
con sid ered *
I f one w ere t o c o n s id e r th e Montana Wo* I com binations ap a rt from th e
o th e r s t o c k - s c io n com b in a tio n s, one m ight q u e s tio n th e v a lu e o f _J* v i r g i n ia n a " as an u n d e rsto c k .
Montana Ho* I u n ite d alm ost tw ic e as w e ll w ith
J* soopulorum as w ith J , v ir g in ia n a *
A p o s s i b l e , but perhaps poor ex p la n a ­
t i o n f o r t h e s e r e s u lt s m ight be in th e f a c t th a t Montana No. I i s a c t u a lly
a v a r ie t y o f J* soopulorum and t h e r e f o r e m ight be exp ected t o do b e t t e r
w ith J , soopulorum than w ith J, v ir g in ia n a .
However, in v ie w o f t h e c l o s e
s i m i l a r i t i e s in s tr u c tu r e and chromosome number o f th e tw o u n derstocks
t h i s h a rd ly seems a v a li d reason f o r t h e d if f e r e n c e s o b serv ed .
The number
o f ta k e s o f b o th Montana No. I com binations are low when compared w ith th o s e
o f L ivid us and Savin and a re perhaps to o low t o be co n sid ered o f any s i g n i ­
f ic a n c e in t h i s in v e s t i g a t i o n .
The low p ercen ta g e ta k e s w ith Montana Ho. I
as compared w it h th e r e l a t i v e l y h ig h p ercen ta g es o f L iv id u s and S avin may
p o s s ib ly b e a t t r ib u t e d t o th e d i f f e r e n t growth h a b its o f Montana Ho. I .
Montana No. I i s u p r ig h t, L ivid us and S avin h o r iz o n t a l.
Any one o f a com bination o f s e v e r a l f a c to r s may b e r e s p o n sib le f o r th e
e x c e e d in g ly poor perform ance o f th e P f i t z e r com b in a tio n s.
I n c o m p a tib ility
may have r e s u lt e d from th e d if f e r e n c e in chromosome number o f th e s c io n
and s t o c k in th e s e co m b in a tio n s,
P f i t g e r i s t e t r a p l o i d , ( l l ) , whereas J e
-
18-
T ir g in ia n a and _£» scopulorum are d ip lo id s (2 9 ) 0 (L iv id u s . S avin and Montana
No. I are d i p l o i d s . )
The f a i l u r e o f th e P f i t z e r com binations might a ls o be
a t t r ib u t e d to th e f a c t th a t th e P f i t z e r s c io n s were a lm o st tw ic e as la r g e
as th e s c io n s o f th e o th e r com b in a tio n s.
E x c e s s iv e t r a n s p ir a t io n by t h e
la r g e P f i t z e r s c io n s may have c r e a te d a w a ter r e la t io n s problem .
And
f i n a l l y , th e P f i t z e r com binations' had a fo u r-d a y s h o r te r p erio d under th e
m is t and warm tem perature than d id th e o th e r co m b in a tio n s.
/
In th e p r e se n t in v e s t ig a t i o n s u c c e s s f u l unions were e s t a b lis h e d f o r
a l l grafts' stu d ie d a n a to m ic a lly in w hich a t l e a s t a p a rt o f th e bark o f
th e s c io n and s t o c k was in c o n ta c t or n e a r ly s o .
In o th e r w ords, as lo n g
as th e bark o f scion" and s to c k was c l o s e enough to perm it th e esta b lish m e n t
o f a c a llu s b r id g e , a cam bial union fo llo w e d and a s u c c e s s f u l union r e s u lt e d .
In a l l but one o f th e u n s u c c e s s fu l g r a f t s , th e bark o f th e s c io n and s t o c k
were so fa r ap a rt t h a t th e c a llu s t i s s u e o f one or b o th h a lv e s o f th e g r a f t
had produced e it h e r s c a r t i s s u e or a periderm b e fo r e a u n ion could be accom­
p lis h e d .
HVhen s c io n and s t o c k a re r e l a t i v e l y n ea r one a n o th er, t h e sequence o f
ev en ts in th e g r a ft un ion c o n s is t s o f ( l ) th e form ation o f a c a llu s b rid g e
(w ith eq u al c o n tr ib u tio n s by s c io n and s to c k ) -follow ed by ( 2) th e d ev elo p ­
ment o f a cam bial b r id g e , and ( 3) th e su bseq u en t p ro d u ctio n o f new secon d ary
xylem and seco n d a ry phloem.
The d evelop m en tal sequence d i f f e r s somewhat,
how ever, in th o s e g r a fts in which s c io n and s to c k are r e l a t i v e l y fa r ap a rt
and th u s , in w hich a r e l a t i v e l y la r g e c a llu s b rid g e i s r e q u ir e d . , H ere,
tr a c h e id s and s ie v e c e l l s d i f f e r e n t i a t e from c a llu s t i s s u e on e it h e r s id e
o f and p r io r t o a c t i v i t y o f th e new ly formed cam bial b r id g e .
This p a tte r n
-
19 -
o f developm ent r e s u lt s in a un ion o f t h e v a s c u la r t i s s u e s o f s c io n and s to c k
p r io r to cam blal a c t i v i t y r a th e r than a fterw a rd , and i s s im ila r t o th e phe­
nomenon rep o rted by Sass (3 5 ) as o ccu rr in g in ap p le when th e r e i s e x c e s s iv e
c a llu s fo rm a tio n .
The le n g th o f tim e req u ired f o r a com plete un ion o f s c io n and s to c k
v a r ie s c o n s id e r a b ly depending upon th e d is ta n c e between s c io n and s to c k ,
and th e d eg ree o f accu racy in align m en t o f t h e t i s s u e s o f th e s c io n and
s to c k .
I t i s apparent t h a t a g r e a te r amount o f c a llu s t i s s u e w i l l be n e c e s ­
sa r y t o f i l l in a la r g e r sp a ce than a sm a lle r one.
L ik ew ise more tim e w i l l
be req u ired fo r oam bial form ation in a w id er b r id g e r o f c a llu s t i s s u e than
in a narrow er one.
The second f a c t o r i s c l e a r ly u n derstood i f one r e c a l l s
th a t a l e s s a c cu ra te a lign m en t o f t i s s u e s o f th e s c io n and s t o c k r e s u lt s in
a g r e a te r r a d ia l s e p a r a tio n o f th e cambia o f s c io n and s t o c k , thus r e q u ir in g
th e form ation o f lo n g e r cam bial b r id g e s than in th o se g r a f t s in which th e
cambia l i e a d ja c e n t t o one a n o th er.
I t i s apparent from th e rev iew o f li t e r a t u r e th a t c o n tro v er sy e x i s t s
w ith regard t o th e r e l a t i v e r o le s p layed by th e cambium and o th er t i s s u e s
i n c a llu s fo rm a tio n ,
tn th e p r e se n t in v e s t ig a t io n th e' cambium appeared t o
c o n tr ib u te l i t t l e , i f a t a l l , t o c a llu s fo rm a tio n .
stu d y were made du rin g w in te r dormancy,
The g r a f t s in t h is
At t h i s tim e o f y e a r th e cam bial
zone o f th e ju n ip e r i s one to th r e e c e l l s w id e.
During t h e same tim e o f
y e a r th e cam bial zones o f o th e r s p e c ie s may p o s se s s as many as ten over­
w in te r in g elem en ts ( 1 6 ) ,
One m ight l o g i c a l l y ex p ect th e cam bial zone o f a
s p e c ie s w ith many o v erw in ter in g elem ents t o p la y a g r e a te r r o le in c a llu s
form ation than one w ith o n ly a few .
I t would be in t e r e s t i n g t o examine
20 «
“»
ju n ip e r m a te r ia l th a t had been g ra fte d d u rin g th e growing sea so n , th a t i s ,
a t a tim e when th e cam b ial zone would c o n s i s t o f more than m erely th e la y e r
o f cam bial i n i t i a l s and one or two d e r i v a t i v e s .
During a p erio d o f cam bial
a c t i v i t y one would ex p ect th e cam bial zone t o c o n tr ib u te much more, to c a llu s
form ation than d u rin g th e dormant p e r io d .
Under any circu m sta n ce, th e r o le
p layed by th e cambium in c a l lu s fo rm a tio n during th e dormant p eriod p robably
v a r ie s from s p e c ie s t o s p e c i e s , in p a rt a t l e a s t , a cco r d in g t o th e number
o f elem en ts o v e r w in te r in g in th e cam bial zo n e.
With regard to th e r e l a t i v e r o le s p layed by o th er t i s s u e s in c a llu s
form ation , p r a c t i c a l l y a l l t i s s u e s o f th e stem have b een c i t e d , a t 'one tim e
or a n o th er, as havin g c o n trib u ted ( s e e rev iew o f l i t e r a t u r e ) .
In th e
p r e se n t in v e s t ig a t i o n i t was observed t h a t th e parenchyma c e l l s o f th e cor=
t e x , phloem, and xylem , in c lu d in g th e phloem and xylem rays., are capable
o f u n d ergoin g c o n s id e r a b le p r o l i f e r a t i o n .
Of t h e s e , t h e parenchyma c e lls -
o f th e xylem and th e xylem rays a re p a r t ic u la r ly im portant when th e t i s s u e s
o f th e bark o f s c io n and s to c k a r e n ot in a lig n m en t.
I t i s in t e r e s t i n g t o
n o te Ifhat, ex cep t f o r immature elem en ts, t h e s e c e l l s a re r a r e ly m entioned
*
in th e l i t e r a t u r e as th e p recu rso rs o f c a llu s t i s s u e .
Very l i k e l y th e
reason f o r t h i s l i e s in th e f a c t t h a t in many s p e c ie s th e parenchyma c e l l s
o f th e xylem and xylem rays p o s s e s s secon d ary w a lls and a r e , t h e r e f o r e , un able
t o undergo r e v e r s ib le changes (ill-).
In o th e r words, i t i s im p o ssib le f o r
th e s e c e l l s to become m e r iste m a tic ,
The parenchyma c e l l s in th e xylem and
xylem rays o f th e ju n ip e r p o s se s s prim ary w a lls and th u s can become m e r is te m a tic ,
S in c e th e p it h was r a r e ly in v o lv e d in t h e u n ion s s tu d ie d , no
d ata was gath ered w ith regard t o i t s r o le in c a llu s fo rm a tio n .
6
-
21 -
The l i t t l e a d d it io n a l eare and tim e req u ired t o be c e r t a in o f align m en t
o f th e two barks would g r e a t ly in c r e a s e th e e f f i c i e n c y o f th e g r a f t in g .
-
22 -
LITEBATtTRE CITED
( 1) A d rian ce, Ge W0, and F0 R0 B r iso n 0 1955« P rop agation o f h o r t ic u l­
t u r a l p l a n t s „ McGraw-Hill Book Co0, I n c oj Hew York0
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growth o f some v a r i e t i e s o f Jiirtiperus, P roc0 Amer0 S o c0 H ort0 S c i ,
5 8 : 3 0 1 -3 0 7 .
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( 7) Chandler, C0 A0' 19)48,
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Amer, Hurs0
( 8 ) Chang, W en-T sai, 1938, S tu d ie s in in c o m p a t ib ilit y betw een s to c k and
s c io n , w it h s p e c ia l r e fe r e n c e t o c e r t a in deciduous f r u i t t r e e s , Jou r,
PomoI , and H ort0 S c i , 15: 267-525«
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on th e i n t e r s p e c i f i c g r a ft u n io n , B d t, Gaz0 19: 5 9 2 -6 0 8 ,
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flo w e r in g p la n t s ,
George A lle n & TJnwin L td ,, London,
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Baume and S tra u ch er0 pp 1=32* C a s s e l0
(1 9 ) Hartmann9 H0 T09 and D0 E0 K e ster0 1959* P la n t p ro p a g a tio n , p r in c ip le s
and p r a c tic e s * P r e n t ic e - H a ll, I n c 0 Englewood C l i f f s , H0 J0
(20) H errero9.. J* 1951» S tu d ie s o f co m p a tib le and in co m p a tib le g r a ft combina
t io n s w ith s p e c i a l r e fe r e n c e t o hardy f r u i t tr e e s *
Jour0 Horb0 S ci*
2 6 ; 186-237»
(2 1 ) H i l l , J 0 B0 1953» Ju n ip er g r a f t in g , p r a c t ic a l and t e c h n ic a l a p sects*
P ro c0 Third Annual Meet* P la n t Prop* Soc* pp 86-93»
( 22) __________ ' ' *
1959»
(2 3 ) Johansen9 D0 A0
In c * , Hew York*
pers* lt r * dtd 13 apr*
I9I+O*
P la n t m icrotech n iq u e*
McGraw-Hill Book Co.*,
( 2)|) J u lia n o 9 J* B* 191+1* C allus developm ent in th e g r a ft.u n io n *
r P h illlp p in e Jour* S ci* 75: 21+5-251+=
(25) K o s to ff9 D0
565- 577.
1928*
(2 6 ) K u ster9 E» 1925»
76-183* Jena0
The
S tu d ie s on c a llu s t is s u e * Amer0 Jour* Bot* 15:
. .........................................................
P a th o lo g isc h e P fla n zen a n a to m ie0
(2 7 ) M ah lsted e,' J* P09 and'E* S 0"Haber*
W iley & Sons, In c * , Hew York*
1957»
3 t e A u fl0
P la n t propagation*
pp
John
(2 8 ) M a llin so n 9 J 0' W0 1926* G raftin g rhododendrons and c h o ic e c o n ife r s*
P art I I I *
Ju n ipers* F l o r i s t s rt Exchange 6 l : 7U9-751»
. ) ‘l j
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(2 9 ) Mehra9 R0 H09 and T1- u , Khoshoo*
Jou r0 Genet* 54$ 165=180*
1956*
C ytology o f c o n ife r s*
I*
(3 0 ) Mossej B09 and "J7 Herrero* 1950* S tu d ie s on in c o m p a t ib ilit y betw een
some p ear and quince g r a fts * Jour* Hort* Sci* 26: 238=21+5»
(3 1 ) O r ffe r 9"C'*" J . '1957* "O ntogenetic and ex p erim en ta l s t u d ie s on th e
e sta b lish m e n t o f th e g r a ft union in th e grapevine* Ph* D* T h e sis9
U* o f C a lif * , Davis*
(3 2 ) P r o e b s tin g 9 'E* L0 1926* S tr u c tu r a l w eaknesses in i n t e r s p e c i f i c
g r a fts o f P y ru s* Bog* Gas0 - 82: 356-338*
( 33)
'
* 1928* F u rth er o b serv a tio n s on s t r u c t u r a l d e f e c t s
o f th e g r a ft u n io n * B o t0 Gaz* 8 6 : 82-92»
(3b) "Roberts, R„ E0
15: k23-k63*
19Wo
T h e o r e tic a l a s p e c ts o f g r a fta g e ,
B ot0 'Rev0
( 55) S ass , J 0 E0
1932« Formation o f c a llu s knots' on a p p le g r a fts as
r e la te d t o th e h is t o lo g y o f th e g r a f t union* B o t„ Gaz0 9W 3&k-380<>
( 56) S h a r p ie s, A*, and H0 Gunnery* 1933« C allus form ation in H ibiscu s
R o sa -sin e n s i s L, and Hevea b r a s i li e n s i s . Mull* Agr. Ann. Bot * tj/f:
8 2 7 -6 3 9 .
™
(3 7 ) S orau er, P0
I92I4.*
Handbi der P fan zen k ran k h eiten 0
( 58) Waugh, F0 A0 - I 90I4..
B u ll0 2: 16 pp.
The g r a ft u n io n ,
(3 9 ) W e lls , J 0 S .
Hew York*
P la n t p ro p a g a tio n p r a c t ic e s .
1957.
(W ) Y erkes, G. E, ■ I9W 0
B u ll* Ho. 1567 Rev.
5"be A u fl0 I 0 Jen a.
Mass0 Agr0 Exp* S t a 0 Tech0
The Macmillan C o.,
P rop agation o f t r e e s and shrubs *
USDA
Farmersr
25 '
J
-
--
.
.
.
.
. PLATEI
Photo grap h ..showing com parative growth o f L ivid u s on J . soopulorum
and £» V ir g in ia n a tt
:
Photograph showing com parative growth o f S avin on J e soopulorum
and _Jtt v ir g in ia n a ,
""
' :
Photograph show ing com parative growth o f Montana'Hoe. I .Calsoi c a lle d
Pocky Mountain Hoe .1} on J e soopulorum and J e v ir g in ia n a , ■
P LA. TB I I
Ae
T ransverse s e c t io n showing o u te r p a r t o f th e xylem ( x ) s th e eam bial
zone C e), and t h e in n e r area o f th e phloem (above t h e eam bial zone) „
The parenb.hyma' .o p ils ' o f .- t h e . rays ( r ) "ppfsess pritiiany... w a lls in b oth
th e phloem ianti..-xyl-ein'e-- The. v e s i c a l ' elem en ts,-,in ''the' .phloem are
f ib e r s ( f ) ^/,parenchyma p ells... (p c )' and 'sieve c e l l s ' , ' ( s c ) e X320e
Be
L on gitu d in al- s e o tio n - shw oing 'o b liq u e Iy P rien ted t r a c h e id s ( t ) th a t
have d if f e r e n t ia t e d : froni' d a llu s t i s s ne ( c a l) b r id g in g s c io n (sn )
and s t o c k ( s t ) . e . XSOe ;,
........
'; ' ;
,
- . •- ■
■■
,
D e t a i l s ; o, c&mbial zone; c a l , c a llu s t i s s u e ; f , f i b e r s p c, parenchyma
c e l l s ; r , rays'; s c , s ie v e c e l l s ; sn , s c io n ; $ t , s to c k ; t ,
t r a c h e id s ; x , xylem*
\
' .v-:
■ :■
- ■■' •.
I
PLATE I I I
A,
,T ransyerse -SBoiiidn' shtiwin'g o u te r p a r t o f the- XyIemi Cx)# th e oamhial
• zorie (■c)y th e /p h lo e m '( p h ) , ; a n i c o r te x ( o t x ) » , The t i s s u e s o f s c io n
■- and .S to ck are' v ery a c c u r a te ly matched ( t h e s to c k i s t o th e -le ft o f
ttie h r q k e r i/lin e , the. s c io n t o th e r i g h t ) . XSO..=
B. • T r a n sv e r se -se c tio n ' showing p a rt o f th e s to c k ( s t ) and s c io n (s n ) ,v
.iriclriding- c a llu s t i s s u e ( c a l ) produced- betw een them from p r o li f e r a t e
. -Irig -xylem- parenchyma and xylem ray parenchyma# X80#
.
■•
C, . T rh nsverSe" se c tio n show ing a U -shaped oam hial un ion betw een s to c k ''
( s t ) a n d 's c io n (sn )# X80#
- v; t •• " • •
.
.
■
.
"fv -.
D etails.?' - c# cam bial zo n e5 c a l , c a l lu s t i s s u e ; o tx , c o r te x ; ph, phloem;-'
s n , s c io n ; s t , s to c k ; x , xylem .
•
. .■
' ;y.
•
'
'
':
•1
- • .,
; KU,
:
•-
/'X:
-v
' : .V
•- L i'
%
1 " ..
31
'
.
•
■.
■PLATE I Y
A 0 ■ T r a n sv e r se ' s e c t i o n -sh ow in g .gap b e tw e e n t h e tw o h a lv e s
. ( s t ) n o t s e p a r a t e d " h y t h e s c i o n ( s n ) » XSO0
o f th e s to c k
B»,'- T r a n s v e r s e " s e c tio n '; sh o w in g ' p a r t o f s t o c k ( s t ) and s c i o n . ( s n ) 0 N o te
t h a t a cambium ( c ) h a s b e e n p ro d u c e d i n t h e c a l l u s t i s s u e l y i n g '
• a d j a c e n t t o b o t h s t o c k and s c i o n ,
XBO0
C0
T r a n s v e r s e s e c t i o n sh o w in g u n io n b e tw e e n s t o c k ( s t ) a n d s c i o n ( s n )
' i n w h ic h t r a o h e i d s ( t ) and s i e v e c e l l s ( s o ) h a v e d i f f e r e n t i a t e d from
c a l l u s t i s s u e on e i t h e r s i d e o f t h e c a m b ia l b r i d g e ( c ) . - XBO0
D0 . T r a n s v e r s e s e c t i o n s h o w in g u n s u c c e s s f u l g r a f t " r e s u l t i n g from t h e
/ S o id n r S-. ( s n ) b e i n g embedded i n t h e s e c o n d a r y x y lem o f t h e s t o c k ( s t ) ,
'./• N o te t h e p erid erm , ( p ) o f t h e s c i o n , ) 'XBO0
"
'
'
D e t a i l s c , c a m b ia l zo n e? p , p e r id e r m ; s c , s i e v e , c e l l s ; S n , s c i o n , s t ,
'
s to c k ; t , tr a o h e id s ,
MONTANA STATE UNIVERSITY LIBRARIES
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