RESPONSE OF CONCORD GRAPE VINE (VITIS LABRUSCA L . ) TO
VARIOUS LEVELS OF ESSENTIAL NUTRIENT ELEMENTS

By

ERNEST L. BERGMAN

AN ABSTRACT

Subm itted to the School for A dvanced Graduate Studies of M ichigan
State U n iv ersity of A g ricu ltu re and Applied S cien ce
in p a rtia l fu lfillm en t of the req u irem en ts
for the d eg ree of
DOCTOR OF PHILOSOPHY
D epartm ent of H orticulture
1958

A pproved

A * ''

//* '

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L

ERNEST L. BERGMAN

ABSTRACT

The r esp o n se of Concord grape v in e s to v a rio u s le v e ls of e s s e n tia l
nutrient e le m e n ts w as studied in the green h ou se w ith th ree ex p erim en ts.
In E xp erim en t I, o n e -y e a r -o ld rooted cuttings w e re u sed .

The

ch eck treatm en t w as supplied with standard Hoagland solution, and high
(5x) or low (Ox) le v e ls of nitrogen, phosphorus, p o ta ssiu m , ca lciu m , m a g ­
nesium , iron, m an ganese, boron, copper, zin c, and m olybdenum w ere o b ­
tained by adjusting the standard solution to the s p e c ific le v e ls .

High le v e ls

of calcium and p otassiu m w ere te ste d in com bination w ith both sulfate and
ch lorid e as anions.

In E xperim en t II, r a tio s of I. 6 to 21. 2 of Ca+Mg
K

the solution w ere u sed to te s t the in flu en ce on grow th and e x p r e ssio n of
p o ta ssiu m d eficien cy .

In E xperim en t III, the e ffe ct of th ree le v e ls of m an ­

g a n ese (2x, 25x and 50x Hoagland solution m anganese) upon plant growth
w as te sted .

In E xp erim en ts II and III, tw o -y e a r -o ld p lan ts w ere u sed and

the standard solution w as m odified by doubling m an ganese and in c r e a sin g
copper by 50%.
T otal lin ea r growth, dry w eight accu m ulation of ro o ts, ste m s, p e tio le s
le a f blad es, and total dry w eight w ere taken.
calcu lated .

The sh o o t/r o o t r a tio s w ere

P e tio le s and ste m s of each plant w ere an alyzed for nitrogen,

phosphorus, p o ta ssiu m , calciu m , m agnesium , iron, m anganese, boron,
copper, zin c, and p a r tia lly for ch lorid e.

ERNEST L. BERGMAN

ABSTRACT - 2

The ch eck treatm en t produced m o st total lin ea r growth, w h ile low
le v e ls of m ajor e le m e n ts, w ith the ex cep tio n o f m agn esium , d e p r e sse d
lin ea r grow th m ore than low le v e ls of m in or e le m e n ts.

The high le v e l of

m olybdenum produced m o st total dry w eight, w h ile low le v e ls of p otassiu m
and nitrogen, and high le v e ls of p o ta ssiu m with ch lo rid e as anion w e re m o st
d e p r e ss iv e on total dry w eight.

T rea tm en ts d eficien t in n itrogen and p h o s­

ph oru s or high in m agn esium and m olybdenum ca u sed the lo w est sh o o t/r o o t
ratio.

High le v e ls of phosphorus and nitrogen, and low le v e ls of m agnesium

c a u sed h ig h est sh o o t/r o o t r a tio s .
A p o sitiv e relation sh ip betw een the s p e c ific nutrient e lem en t in the
solu tion and in the p e tio le s w as o b serv ed for a ll e lem en ts ex cep t for iron
and zin c.

P e tio le s see m e d to provide a b etter indication of norm al or above

norm al le v e ls of nutrition than the ste m s.
indicate b etter low or d eficien t le v e ls .

The ste m s, how ever, see m e d to

The stem content w as alw ays equal

to, or h igh er than, the p e tio le content under low or d eficien t nutrient co n d i­
tio n s.
The Ca+Mg r a tio s in the nutrient solution w ere of no d ir e ct influence
K
on grow th o r on c o lo r of p otassiu m d eficien cy sym p tom s.

Indirectly, how ­

e v er , p o ta ssiu m , calciu m , and m agn esiu m e x e r te d v a ried in flu en ces.
C hronological appearance o f v is ib le d e fic ie n c y or to x ic ity sym ptom s

ABSTRACT - 3

ERNEST L. BERGMAN

w ere not c o r r e la te d w ith the d e p r e ssio n of grow th r e su ltin g from the sp e c ific
elem en t.

A ll low le v e l and four of the high le v e l trea tm en ts produced v is ib le

lea f sym p tom s.

M agnesium d eficien cy sym ptom s on le a v e s developed m o re

rapidly w ith high p o ta ssiu m than when m agn esium w as om itted.

V isib le d e ­

fic ie n c y sym ptom s on plan ts coincided w ith the follow ing v a lu e s in p e tio les:
n itrogen . 51%, phosphorus .10%, p otassiu m .47%, calcium .26%, m agnesium
. 14%, m an gan ese 18 ppm, boron 14 ppm, copper 28 ppm, and zin c 11 ppm.
Many sign ifican t in tera ctio n s betw een nutrient e lem en ts w ere found.
G en era lly m ajor e lem en ts w ere of m o re influence on the m in or elem en ts and
v ic e v e r s a .

High le v e ls of m ajor e lem en ts in the nutrient solution had m o re

influence on nutrient absorption than high le v e ls of m inor e le m e n ts.

Some

of the m o re strik in g in teraction s betw een elem en ts were a s fo llo w s: (1) Low
n itrogen in the solution ca u sed low phosphorus, low boron, and high calcium
in the p e tio le s .

(2) High nitrogen produced high phosphorus and high boron

p e tio le content.

(3) Low phosphorus in p e tio le s coin cid ed w ith high m angan­

e s e and high iron.

(4) V ery low le v e ls of copper and zin c w ere found under

d eficien t p o ta ssiu m conditions.

(5) A sp e c ific amount of p o ta ssiu m appeared

to be n e c e s s a r y for adequate absorption o f calcium and m an gan ese.

(6) M ore

p otassiu m w a s absorbed w ith sulfate than w ith ch lo rid e a s an anion.

In the

la tter c a se , m ore calciu m w as found in p e tio le s .

(7) High nitrogen, p h o s­

phorus, p otassiu m , m anganese, and boron in p e tio le s appeared under calcium

ERNEST L. BERGMAN

d eficien t con d ition s.

ABSTRACT - 4

(8) A sign ifican t p o sitiv e c o r r e la tio n betw een total iron

and m an ganese in the p e tio le s w as e sta b lish e d .

(9) Low phosphorus, p o ta s ­

sium , calciu m , and high m agnesium w ere a s so c ia te d w ith high m an ganese
v a lu e s in p e tio le s .

(10) M ore boron w as o b serv ed in p e tio le s of plan ts trea ted

w ith high le v e ls of nitrogen, phosphorus, p o ta ssiu m , and low le v e ls of c a l­
cium than in th ose plan ts treated with high le v e ls of boron.
With e x tr e m e ly high m anganese con cen tration s in the solution, m an­
g a n ese to x ic ity sym ptom s appeared on le a v e s .

Growth w as not a ffected at

th is le v e l, how ever, the b e r r ie s did not ripen ev en ly within c lu s te r s .

RESPONSE OF CONCORD GRAPE VINE (VITIS LABRUSCA L .) TO
VARIOUS LEVELS OF ESSENTIAL NUTRIENT ELEM ENTS

By

ERNEST L. BERGMAN

A THESIS
Subm itted to the School for Advanced Graduate Studies of M ichigan
State U n iv ersity of A gricu ltu re and A pplied S cien ce
in p artial fu lfillm en t of the req u irem en ts
for the d eg ree of

DOCTOR OF PHILOSOPHY
D epartm ent of H orticulture

1958

ProQuest Number: 10008588

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ACKNOWLEDGEMENTS

The author w ish e s to ex p ress his sin cere thanks and appreciation
to Dr. A. L. Kenworthy for his a ssista n ce in outlining the problem , ca rry ­
ing out the experim ental work, and preparing the m anuscript; to Dr s. R. L.
Carolus, G. P. Steinbauer, R. L. Cook, and K. Lawton for their guidance
and editing of the m anuscript; to Dr. E. J. Benne and Mr. S. Bass and
th eir staff for carrying out the chem ical analyses; to Dr. J. D. Downes
for his help on certain a sp ects of sta tistica l an alyses, and to Dr s. H. K.
Bell and R. P. L arsen for their technical a ssista n ce.
Special recognition is given to Dr. E. J. Kraus (visitin g p rofessor,
Department of H orticulture, Oregon State College, C orvallis, Oregon,) for
encouraging the author to per sue further education.
Special acknowledgement is a lso made to the author's wife, A lice,
for her u n selfish sa c r ific e s, constant a ssista n ce and encouragem ent through­
out the years, and to the National Grape Cooperative A ssociation for p ro ­
viding the financial grant for carrying out the research .

TABLE OF CONTENTS

Page
INTRODUCTION

.............................................

.

1

REVIEW OF LITERATURE

......................................................

2

EXPERIMENTAL PROCEDURE. .

.......................................................

21

EXPERIMENT I. EFFEC T OF VARIOUS LEVELS OF 11 NUTRIENT
ELEMENTS ON GROWTH, PETIOLE AND STEM COMPOSI­
TION . . .
......................... . . . . . . . . . . . . . .
22
RESULTS: DEFICIENCY AND TOXICITY SYMPTOMS OF STEMS
AND LEAVES . . . . . .
. . . . . . . . . . . . .
. .
25
VISUAL SYMPTOMS ON ROOTS

......................

43

GROWTH. . . . . . . . . . . . . . . . . . .

46

NUTRIENT ELEMENT COMPOSITION OF PETIOLES
AND STEMS.
......................

54

DISCUSSION. . . . . . . .

. . . . . . . .

. . . . . .

72

EXPERIMENT II. EFFEC T OF Ca^Mg RATIOS ON GROWTH,
PETIOLE AND STEM COMPOSITION . . . . . . . . . .

86

EXPERIMENTAL PROCEDURE. . . . . . . . . . . . . .

86

RESULTS: GENERAL OBSERVATIONS . . . .

88

. . . . . .

VISUAL APPEARANCE OF ROOTS . . . . . . .
GROWTH. . . . . . . . .

. . . . . . . . .

ELEMENTAL COMPOSITION OF PETIOLES AND
STEMS. . . . . . . . . . . . . . . . . .

89
.

90

91

CONTENTS CONT'D
Page
DISCUSSION.........................................................................................
EXPERIMENT III. E F F E C T OF THREE LEVELS OF MANGAN­
ESE ON GROWTH, PETIOLE AND STEM COMPOSITION. .

105

I ll

EXPERIMENTAL PROCEDURE....................................................

I ll

RESULTS: GENERAL OBSERVATIO NS..................................

112

VISUAL SYMPTOMS.........................................................

112

GROWTH................................................................................

114

ELEM ENTAL COMPOSITION OF PETIOLES AND
STEMS...........................................................................

116

DISCUSSION.........................................................................................

118

SUMMARY............................................................................................................

121

LITERATURE C IT E D ....................................................................................

126

APPENDIX

134

INTRODUCTION

M ichigan ranks third in the United States in the production of the
Concord grape (V itis labrusca).

The incom e from grapes provides a large

share of the farm incom e receiv ed by many fruit grow ers in the South­
w estern counties of M ichigan.

The nutritional status of M ichigan grape

vineyards im proved considerably after it was found that potassium was
lim itin g grape production (Larsen, 1955).
In extrem e c a se s, deficiency or toxicity sym ptom s may occur on
le a v e s.

If the shortage is not critical, the plant may show reduced growth

an d /or production.

A visu al diagnosis in this instance is very difficult.

Petiole an alyses have opened new avenues for interpretation and c o r r e c t­
ion of nutritional d isord ers p rior to the appearance of visual sym ptom s.
The purpose of the p resent investigation was to study the respon se
of Concord grapes to various le v e ls of essen tia l nutrient elem ents, as
shown by visu al lea f sym ptom s, growth, p etiole and stem com position.
The data indicate the behavior of certain elem ents within the plant and
their interactions with other elem en ts.

They also suggest p o ssib le c o r ­

rection of visu al or hidden nutritional disord ers in Concord grapes.

2.

REVIEW OF LITERATURE

The influence of nitrogen (N), ph osphorus (P), p o ta ssiu m (K), c a l­
cium (Ca), m agn esium (Mg), iron (F e), m an gan ese (Mn), boron (B), copper
(Cu), zin c (Zn), sulphur (S), ch lorin e (Cl), and m olybdenum (Mo) upon
grow th and production of g ra p es has been the subject of num erous in v e s ti­
ga tion s.

The id en tification of c er ta in d e ficien cy or to x ic ity sym ptom s

w ith the e lem en ta l com p osition of a c er ta in plant part has been fu rthered
in r ec e n t y e a r s .

Lagatu (1927), after finding r e s p o n se s to N, P and K,

applied in f e r tiliz e r by m ean s of le a f a n a ly sis, w rote
"The d ia g n o sis b ased on the ch em ica l com p osition of the
le a f is a s a ccu rate and in d ica tiv e for the in ten sity a s it
is for the nature of nutritional supply".
Since then, m any d ifferen t w ays of e x p r e s s in g the m o st su itab le
e lem en ta l com p osition for a sa tisfa c to r y production and a healthy plant
have been p r e se n ted .

Lagatu and M aume (1940) fe lt that the ratio of

FhP^O :K O in the le a v e s should be 41:8:51, w h erea s V ettori (1954) thought
45. 5:9:45. 5 w as the b est, and L iveran t(1955) su g g ested a ratio of 64. 1:11. 6 :
24. 3.

Vidal (1955) added th ese th ree e lem en ts togeth er and rep o rted 3. 9%

dry w eight a s a v era g e for good vin ey a rd s w ith an optimum of 4. 33%.
G oodall and G regory (1947) r ev ie w e d the v a rio u s r ep o rts and concluded
that no v in e s resp on d ed to fe r tiliz e r when the p er cen t dry w eight content

3

of th eir le a v e s in fa ll w as above 1 .5 N, . 19 P, and 1. 03 K.
Beattie and F o r sh e y (1954) o b se rv e d in Ohio that a C oncord grape
p e tio le co m p osition (per cen t dry w eight) of . 77 N, . 14 P, 2 K, . 7 Ca, and
. 15 M g w as n e c e s s a r y to produce 3. 5 tons of g ra p es p er a c r e .

T h e se v a lu es

a r e s im ila r to L a r s e n ’s (1957) standard v a lu e s (per cent dry w eight) in
p e tio le s for M ichigan, w hich a re . 8 N, . 24 P, 1. 91 K, 1. 60 Ca, and . 46 Mg.
Other v a lu e s m entioned by L a rsen w ere 44 ppm Fe, 434 ppm Mn, 25 ppm
B, 35 ppm Cu, and 23 ppm Zn.

Coombe and A llan (1955) conducted a n u tri­

tional study of v in ifera type grap es grown in deep w hite sand in A u stra lia
and found the follow in g v a lu e s (per cent dry w eight) for le a v e s of v ig o ro u s
plants: 2. 55 N, . 52 P2 0 ;y !• 48 K^O, 3. 08 CaO, and . 76 MgO.
M agoon et al. (1938), in a sp ectro g ra p h ic a n a ly sis, com pared Con­
cord with the Ontario v a r ie ty and found the follow in g r e s p e c tiv e v a lu e s in
p e tio le s:

. 61 and . 76% P, 1. 97 and 2. 20% K, . 57 and . 47% Ca, . 28 and

. 22% Mg, 190 and 220 ppm F e, 180 and 220 ppm Mn, 17 and 16 ppm B,
41 and 47 ppm Cu, 450 and 410 ppm A l.

V idal (1955) drew attention to the

fact that in M aroccan v in ife ra vin ey a rd s the nitrogen and ph osphorus content
in le a v e s w as h ig h est in 3- and 4 -y e a r -o ld plan tation s, and lo w e st in th o se
of over 27 y e a r s old.

He a r r iv e d at a com bined value of 4. 62% for young

v in ey a rd s when N, P2O5 and K^O content w ere added, and 3. 69% for o ld er
on es, c le a r ly ind icating that w ith in c r e a sin g age, nitrogen and phosphorus
b ecam e d eficien t.

4

N itrogen:

The a s so c ia tio n of a d efin ite amount of nitrogen in the

plant w ith v isu a l sym p tom s and w ithout sym p tom s is not too c le a r b eca u se
of ra n g es in co m p o sitio n and se a so n of sam plin g.

Shaulis and K im ball in

New Y ork (1956) su g g ested that a Concord grape p e tio le content o f below
1. 5% N in June a s approaching a d eficien cy , and found a range of 1. 7 to
2.4% N for v igorou s y a rd s.

They a ls o rep o rted a sea so n a l d e c r e a s e of

nitrogen w ithin the plant, w hich a g reed w ith Lagatu" s (1927) fin din gs.
W agner (1924) rep orted 1. 0 to 2. 0% n itrogen in w hole le a v e s and thought
that a m ature le a f should contain about 1. 55% N.

M aume and Dulac (1948)

rep o rted 2. 6% N in le a v e s at the beginning of the sea so n , and 1. 6% N at
m atu rity a s norm al for v in ifera type g ra p es in F ran ce.

M cG eorge and

B reazeale (1955) found a d e c r e a s e from 2. 3% N in A pril to . 96% in le a v e s
of T hom pson s e e d le s s in A rizona, in July.

Bergm an and Kenworthy (1957)

e sta b lish e d the sam e trend in Concord grape p e tio le content from 1. 34% N
in June to . 68% N in Septem ber, without appearance of v isu a l N d e ficien cy
sym p tom s.
H e rsc h le r (1933), w ith pot ex p erim en ts, a s w e ll a s H agler and Scott
(1949), w orking with sand cultu re, found le a v e s turning y e llo w ish g reen
a fter a few w eek s of growth without n itrogen .

Shaulis and K im ball (1956)

a s so c ia te d short internode length and stoppage of lin ea r grow th by m id-July
w ith nitrogen d e fic ie n c y .

W hereas, Coombe and A llan (1955) found a higher

5.

n itrogen content in le a v e s of stunted p lan ts (2. 87%) than in v ig o r o u sly g r o w ­
ing on es (2. 55%), K obayashi et a l (1955) found 2. 28 to 2. 75% N in le a v e s of
n o n -b ea rin g and 2. 23 to 2. 61% N in b ea rin g p lan ts of the D elaw are v a r iety .
P artridge and V eatch (1931), a s w ell a s C ooper and V aile (1939) o b s e r ­
ved m ore vig o ro u s grow th and h igh er y ie ld s w h ere n itrogen f e r t iliz e r s w ere
applied.

U lrich (1942a) rep orted an in c r e a s e in nitrogen content of p e tio le s

by applying one pound of (N H ^ S O ^ p er vin e for th ree y e a r s .

P e tio le s of

untreated p lan ts show ed 1. 09% N in May and . 47% N in Septem ber, a s c o m ­
p ared to 1. 49 and . 62% N r e s p e c tiv e ly of trea ted p lan ts.

L ea v es from the

sam e un treated p lo ts had 3. 48 and 2. 21% N, w h e r ea s th o se from trea ted
p lo ts had 3. 95 and 2. 49% N.
V olk (1938) ca lcu la ted the ratio of root to above-ground p a r ts of p lan ts
grow n for two y e a r s in w ater cu ltu re and e sta b lish e d the fo llo w in g r o o t/sh o o t
ra tio s:

for m inus N 1:1.81; for high N 1:4. 56, and for the check 1:3. 6.

The

m in us N plan ts had an e x c e lle n t r o o tsto ck w ith ligh t yellow to yellow brown
c o lo r and m ore wood than le a v es; the high nitrogen p lan ts, how ever, p r o ­
duced m any m ore le a v e s .
P h osp horus:

V ery few o b serv a tio n s of phosphorus d e fic ie n c y under

fie ld conditions have been rep orted .

C ais (1954) found such a fie ld in Italy

and o b se rv e d v e ry sh ort growth, sm a ll te r m in a ls, ch lo r o tic le a v e s w ith
o c ca sio n a l m argin al d e ssic a tio n and flo w er drop.

If, how ever, pollin ation

6.

had taken p la ce, the b e r r ie s often d e s sic a te d but sta y ed on the vine, looking
lik e "pepper g r a in s ” .

T h is condition w as c le a r e d up by the ap p lication of

p h osphorus f e r t iliz e r s alone, w h erea s other f e r t iliz e r s w ere o f no influence.
A sim ila r field w as o b serv ed by S tellw aag (1955) near M eran, Italy.

T his

fie ld w as strik in g by the dark g reen appearance of the le a v e s and v e ry short
te r m in a l growth.
Lagatu and M aume (1927), and P artridge and V eatch (1931) o b serv ed
so m e r esp o n se to ph osphorus fe r tiliz e r s , but fe lt the r esp o n se w as due to a
b etter balance betw een N, P, and K rather than due to phosphorus alone.
C ooper and V a ile (1939), a s w e ll a s Randolph (1944), rep o rted r e sp o n se of
phosphorus on growth and y ield of C arm en g ra p es in T ex a s.

H e r sc h le r

(1933), on the other hand, w as not able to c o r r e c t s e v e r e phosphorus d e fi­
c ie n c y sum ptom s with ph osphorus.
having . 22% P content.

The sym ptom s o c cu rr e d on le a v e s

W agner (1924) fe lt that . 09% P in the le a f w as su f­

fic ie n t inasm uch a s th ere o c cu rr e d a w ide range under fie ld conditions.
M aume and D ulac (1948) rep o rted v a lu es in le a v e s v aryin g from . 1% P
at the beginning of the se a so n to . 06% P at the end of the sea so n .

A rch ibald

(1956) rep orted a sea so n a l d e c r e a se from . 32 to .11% P, and Bergman and
Kenworthy (1957) found a sea so n a l change from . 27 to . 21% P in p e tio le s .
F o r sh e y (1955) o b serv ed that phosphorus is m o re constant in p e tio le s than in
le a f blad es throughout the grow ing sea so n .

7.

Volk (1938), w orking with w a ter cu ltu re, e sta b lish e d the follow in g
r o o t/sh o o t ratios:

1:3. 03 for m inus P trea tm en ts; 1:4. 1 for high P tr e a t­

m ents; and 1 :3 .6 for the check.
P o ta ssiu m :

Babo and M ack (1910), in th eir book on grape g row in g,

d e scr ib e d sym p tom s w hich a r e known today a s being p o ta ssiu m d e fic ie n c y
under the nam e of "Blattbraeune", "B raunfleckigkeit”, and " B runissure”.
W ilhelm (1950) d e sc r ib e d th ree d ifferen t p otassiu m d e fic ie n c y sta g es;
stage 1 (e a s ie s t to c o r r e c t) show ed purpling on the upper su rfa ce (tow ards
the light) o f the le a f at the beginning of May, follow ed by browning from
the m argin s and in terv ein a l n e c r o s is by June.

Stage 2 exhibited purple

c o lo rin g v ery ea rly in the sea so n w ith w eakening o f v in e s and uneven r ip e n ­
ing of the b e r r ie s .

By the end of July, the b asal le a v e s w e r e n ecro tic, but the

tip le a v e s w ere not affected .

In stage 3, the flo w e r s died and the v in e s show ed

a v e r y poor y ie ld as w ell a s v e r y w eak growth.
p ale green in stea d of dark green and b lu e -v io le t.

In th is sta g e the le a v e s w ere
G ollm ick (1955) fe lt that

th ese p otassiu m d e fic ie n c y sym ptom s w e re brought about by the "idle ru n ­
ning" of the a s sim ila tio n apparatus which, due to ph oto-oxyd ative d estru ctio n
of chlorophyll, produced n e c r o s is of c e ll s .
sym ptom s on C oncord grape le a v e s in Ohio.

Beattie (1955) found two d istin ct
W hereas one type show ed p u r ­

p lish -b la c k a r e a s betw een the v e in s, w hich developed into a black leaf, the
other type exh ib ited only m arginal sco r c h with la te r death of the tis s u e .

L a rse n (1955) o b se rv e d the sa m e sym p tom s in M ichigan v in ey a rd s.
H agler and Scott (1953) developed s im ila r sym p tom s on M uscadine
plan ts in sand cu ltu re after four w eek s of growth.

H arding (1957) tr ie d to

rep rod uce the above d e scr ib e d d eficien cy sym ptom s in nutrient cultu re on
C oncord grape v in e s, and w as able to get only so m e purpling of the le a v e s .
U lrich (1942b) rep orted 2. 58% K in May, and . 31% K in Septem ber
in p e tio le s from un treated plan ts, w h erea s plan ts trea ted for th ree y e a r s
w ith p o ta ssiu m show ed 3. 59 and . 8% K r e s p e c tiv e ly .

L eaf b lad es of the

sa m e p lan ts show ed a d e c r e a s e from 1. 02 to . 44% K in untreated p lan ts,
and from 1. 22 to . 57% K in trea ted plan ts.

T his sea so n a l d e c r e a s e of p o t­

a ssiu m w as a ls o rep orted by M cG eorge and B reazeale (1955), Shaulis and
K im ball (1956), A rchibald (1956), and Bergman and Kenworthy (1957).
L a rsen (1955) a s so c ia te d 1. 0% K in p e tio le s in late July w ith a crop
o f below four tons, and a p e tio le content of above 2. 0% K w ith a crop of o v er
s ix ton s.

A rchibald (1956) a s so c ia te d p e tio le content and crop a s follow s:

. 88% K w ith four tons and over; . 47% K with th ree to th ree and o n e -h a lf tons;
and . 24% K with below two ton s.

Shaulis and Kim ball (1956) found in v ig o r ­

ous vin eyard s a wide range of p otassiu m (. 4 to 2. 0%) in p e tio le s , but su g ­
g e sted that the p lan ts becam e d eficien t if the p e tio le content w as below . 6% K
in late A ugust.

Bergm an and Kenworthy (1957) found black le a v e s in two v in e ­

yard s during la te A ugust, one had a p e tio le content of . 44% and the other . 97%

9.

K obayashi et al. (1955) rep o rted that w ith in c r e a s in g p o ta ssiu m in
le a v e s , the ratio of fru it to le a f is a lso in crea sed ; how ever, beyond a c er ta in
le v e l of p o ta ssiu m , fru it yield and amount of fo lia g e d e c r e a se d again.

T hey

found an a v era g e of 2. 1% K in le a v e s of b earin g D elaw are v in es a s com pared
to 1 .5 to 1. 73% K in le a v e s of n on -b ea rin g p la n ts.

Stunted G renache g ra p es

in A u stralia, a cco rd in g to Coom be and A llan (1955), had . 68%
le a v e s and v ig o ro u s p lan ts 1. 48% K^O.

in the

They a lso o b se rv e d le a v e s w ith m a r ­

ginal sco r c h in g and fe lt that . 3% K^O w as a fa ir ly low le v e l.

Boynton (1945)

found s e v e r e p otassiu m d eficien cy sym ptom s in Septem ber on Portland and
D elaw are le a v e s having .25% K content, but only slig h t sym ptom s on the on es
having .61% K at the sam e tim e.
C alcium :

No r ep o rts of calcium d e ficien cy sym ptom s in v in ey a rd s

have been found, and the sym ptom s d escrib ed w ere a ll produced in nutrient
solu tion cu ltu re s.

H agler and Scott (1953) produced calcium d eficien cy sy m p ­

to m s in M uscadine grape v in e s after eight w eek s of grow th by u sin g a calcium
d eficien t solu tion .

Young le a v e s exhibited in terv ein a l and m a rg in a l c h lo r o sis

follow ed by pinhead sp o ts near the m argin of the leaf, d ie -b a ck of the tip and
drop -off.

S tellw aag and Knickmann (1955) d e scr ib e d sim ila r sym p tom s and

m entioned a "beaklike" cu rlin g of the young le a v e s .
S arosi (1956) an alyzed le a v e s and ro o ts of ch lo r o tic v in e s in Hungary,
w hich r ea c te d favorably to calcium a ceta te trea tm en ts.

He found in abnorm al

10 .

le a v e s (s m a lle r and thinner than norm al) m o re iron, p o ta ssiu m , and sodium
but le s s calciu m than in healthy le a v e s .

In oth er w ord s, m onovalent ca tio n s

in c r e a s e d at the e x p en se of divalent.
On the other hand,
c ien cy .

high le v e ls of calcium m ay induce p o ta ssiu m d e fi­

H arding (1957) induced c h lo r o sis on C oncord grape le a v e s in sand

cu ltu re by adding high le v e ls of calciu m when p o ta ssiu m w as d eleted and
p reven ted c h lo r o sis when 4 m ili-e q u iv a le n ts of p o ta ssiu m w ere added.
Boynton (1945) o b serv ed 2. 71% Ca in Portland, and 1. 99% Ca in D elaw are
le a v e s when p otassiu m w as deficien t.
In crea sin g le v e ls of calcium w ith the advance of the se a so n have
been rep o rted by W ebster and C r o ss (1942).

C oncord le a v e s in c r e a se d from

. 45% Ca in m id-June to 1. 05%, and W orden le a v e s from . 48% to . 85%.
Bergm an and Kenworthy (1957) found a sea so n a l in c r e a s e of calciu m in Con­
cord grape p e tio le s from 1. 0% in June to 1. 6% in Septem ber.
M agn esiu m :

Scott and Scott (1952) drew the attention to d iffe re n c e s

betw een v a r ie tie s in the developm ent o f m agnesium d e fic ie n c y sym p tom s and
in th eir r e sp o n se to spray or so il application trea tm en ts.

T hey o b se rv e d that

when no r e sp o n se to m agn esium application occu rred , no in c r e a s e in le a f m a g ­
nesium took p lace, and a s so c ia te d c h lo r o sis with le v e ls below . 18% Mg in
le a v e s .

B eyers (1955) d e scr ib e d d e fic ie n c y sym ptom s as a ch lo ro tic pattern,

extendin g betw een m ain v ein s, w ell defined w ith broad z o n e s of p a le g reen

11.

a lo n g the m argin of the le a f.

The c h lo ro tic p ale g reen le a f tis s u e turned

y ellow in Waltham C r o ss (white grape), and red in Berlinka (blue grape).
O lder le a v e s , at the b ase of the p lan ts, w ere fir s t a ffected .

On Jam es and

Scuppernong (V itis rotun difolia), Lot (1948) o b se rv e d in A ugust that basal
le a f tis s u e m ost rem ote from v e in s and m argin turned p a le g r e e n /y e llo w
w ith the p rim a ry v e in s stayin g green .

He a s so c ia te d s e v e r e sym ptom s on

Jam es with . 09% M g and on Scuppernong w ith . 06% Mg.
crib ed two d istin ct kinds of le a f sym p tom s.

GMrtel (1957) d e s ­

One o ccu rred in sp rin g when

the ligh t g reen in terv ein a l c h lo r o sis turned to red d ish brown on P ortu guese
g ra p es (red grape), follow ed by dropping of the lea f.
only in fa ll with yellow in tervein al c h lo r o sis.
grow th interrupted.

The other o ccu rred

In n eith er c a se w as lin ea r

G erb er and Bussm ann (1955) in Sw itzerland found that

p rem a tu re fall c o lo rin g of le a v e s w as co n tro lled by se v e r a l ap p lication s o f
2. 0% MgO or MgSO^ sp rays w here no r esp o n se had been found to Hoagland
tr a c e elem en t solution or MgSO^ so il app lication.
sa tisfa c to r y in th is c a s e than MgSO

4

MgO sp ray w a s m ore

due to its r e s is ta n c e to rain w ashing.

L a rsen et aL (1957) rep orted no sig n ifica n t in c r e a se of m agn esium in p e tio le s
of C oncord grap es tr e a ted for fiv e y e a r s by MgSO^ so il app lication.
H agler and Scott (1949) produced d e ficien cy sym p tom s in sand cu ltu re
on M uscadine g r a p e s.

Intervein al c h lo r o sis appeared a fter four w eek s on

o ld er le a v e s on the m id -s e c tio n of the vine.

T h ese le a v e s dropped off la te r .

12.

S tellw aag (1954) o b se r v e d that m agn esium d e fic ie n c y sym p tom s could be
found on p lan ts w ith or without m agnesium p r e se n t.

T h is phenom ena m ore

or l e s s depended upon the com p ositio n of nutrient solution, e. g. e x c e s s iv e
am ounts of N and K w ould ren d er a ssim ila tio n o f m agn esium d ifficu lt.
K enworthy (1957) found m agnesium d e fic ie n c y sym p tom s on Concord
le a v e s w h ere the p e tio le content w as . 09% Mg.

P e tio le s of healthy le a v e s

w ithin the sam e vin eyard had . 21% M g content.

G&rtel (1955) ca lcu la ted a

d efin ite ratio betw een

and Mg in R ie slin g grape le a v e s .

H ealthy le a v e s

had a content of 3. 91% K^O and . 08% Mg, o r a tratio of 48. 87.
w eak sym p tom s contained 4. 28%
He su g g ested that the

V in es with

and . 004% Mg, or a ratio of 600. 57.

ratio w ith p o ta ssiu m d e ficien cy w as below 1,

w h e r ea s w ith m agn esium d e fic ie n c y the ratio w as 30 - 100.

B eyers (1955)

o b se rv e d in South A frica that v in es trea ted with (NH^^SO^ had an in c r e a se d
m agn esiu m and red uced p otassiu m uptake, w h erea s in p o ta ssiu m trea ted
p lo ts, d iso r d e r s due to p otassiu m induced m agn esium d e ficien cy appeared.
L a rsen (1957), how ever, found no tr a c e s of m agn esium d e ficien cy sym ptom s
on Concord v in e s w here 1, 000 pounds o f

p er a c r e w ere applied 3 y e a r s

in su c c e s sio n .
Iron:

Iron c h lo r o s is in New M exico w a s rep orted on g ra p es by

C raw ford (1939).

Wann (1941) noted that the high lim e content o f the Utah

s o il w as in ter fe r in g w ith the norm al u tiliza tio n of iron.

The c h lo r o s is v a ried

13.

from y ea r to year, but the dam age w as m o st s e v e r e during m id -su m m e r
p e r io d s of high ligh t in ten sity and high te m p er a tu r e s.

S a m ish (1954) ob ­

s e r v e d in I sr a e l that lim e-in d u ce d c h lo r o s is ca u sed la te foliation , red uced
shoot and length growth, poor germ in ation of p ollen , and th ereb y d eveloped
"shot" b e r r ie s (m illeran d age) on M adeleine O berlin g r a p e s.

In th is c a se ,

le a f ch lorop h yll content, d egree of p o llen germ ination , and total shoot grow th
w e r e d ir e c tly rela ted to the amount of a ctiv e and total iron w ithin the plant.
M anganese:

M aum e and D ulac (1952) rep o rted m ore m an ganese

in young than in old le a v e s .

The g r e a te st variation , how ever, w as found to

be due to location and s o il.

The sam e v a r ie ty v a ried from 2 to 14 m g /1 0 0

g ra m s dry m atter of le a v e s in one location to 1. 5 to 200 m g in another.
Through a n a ly sis, G hrtel (1956) esta b lish e d that le a v e s and p e tio le s w ere
high est, b erry skin and se e d s low est, and wood in term ed ia te in m an gan ese
content.

He a lso found that the m an ganese content w as s p e c ific to s o il and

not to v a r iety .

Beattie and F o r sh e y (1954) o b se rv e d that m an gan ese d e fi­

c ie n c y can be exp ected to o ccu r when the p e tio le content of C oncord gra p es
in July fa lls below 30 ppm .

Bergm an and Kenworthy (1957) rep o rted a 5 -fo ld

in c r e a s e of m an gan ese during the grow ing se a so n in fie ld s having both high
(from 540 ppm to 2, 700 ppm) and low le v e ls (from 68 ppm to 320 ppm ).
L bhm is (1951) in c r e a se d the m an gan ese content in grape le a v e s grown
from cuttings from 620 ppm to 865 ppm when the m an gan ese content of the

14.

solu tion w as changed from 12. 5 m g M n /lite r to 50 m g M n /lite r .
B eyers (1955) o b se r v e d an accentuation of m an gan ese uptake through
am m onium su lfate s o il app lication.

Am m onium su lfa te applied at a rate of

800 pounds p er a c r e in c r e a se d nitrogen, m agn esium and m an gan ese in le a v e s
from 1. 62% N, . 17% Mg, and 366 ppm Mn to 1. 81% N, .19% Mg, and 646 ppm
Mn, w h ile p o ta ssiu m w as d e c r e a se d from . 92 to .83%.
Beattie (1955) rep orted both m an ganese and p o ta ssiu m d e fic ie n c y
sym p tom s on le a v e s w ithin the sam e Concord grape vineyard, but seldom
found typ ical v isu a l sym p tom s of both.

M anganese d eficien cy appeared fir s t

during June-July on the b asal le a v e s of current se a so n shoot growth, and w as
in d icated by the a b sen ce of g reen co lo rin g m a tter in a r e a s betw een v ein s
w hich turned into light yellow green with dark g reen vein ation .

Kenworthy

(1957) found sim ila r sym ptom s in a M ichigan C oncord grape vin eyard .
c a tio n s of KC1 and K^SO^ did not c o r r e c t p o ta ssiu m d eficien cy .

A p p li­

W hile p e tio le s

of untreated v in es show ed .41% K and 11 ppm Mn, v in e s trea ted w ith 300 pounds
KC1 show ed . 47% K and 18 ppm Mn, and v in e s trea ted w ith 360 pounds
had a content of . 34% K and 14 ppm Mn.
d u P le s s is (1948) su g g ested that m ottlin g of grape le a v e s in South A frica
m ight be due to e x c e s s m an gan ese.
ses.

H ow ever, he did not rep ort any a n a ly ­

15.

Boron:

A ccord in g to Branas (1954), a s e a r ly a s 1894, L eC ocq

d e scr ib e d a d is e a s e c a lle d "la m arom ba" on g ra p es in Portugal, which
w as la te r id en tified a s boron d e fic ie n c y by D ias (1953).

Ono et al. (1956)

rep orted from Japan that 20 to 40 pounds of b o ric a cid p er a cre, or . 3 to
. 5% sp ra y s im p roved growth, fru it set, bunch w eight and y ie ld on g ra p es
having "E bi” d is e a s e .
Scott (1941) found great d iffe r e n c e s in v a r ieta l r esp o n se to boron.
Ontario, C arm en and D elaw are v a r ie tie s showed e a r ly sea so n dw arfing
and ex trem e la te r a l bud grow th with no fruit production.

Concord, Worden

and N iagara w ere only m o d era tely affected and exhibited ch lo ro tic pattern
on le a v e s, little stunting and low fruit production.
and Portland show ed no apparent d eficien cy .

G olden M uscat, F redonia

Catawba le a v e s with d e fic ie n c y

sym p tom s contained 6 ppm B, as com pared to 18 ppm B in healthy le a v e s .

A

s im ila r com p arison show ed 6 ppm and 23 ppm for le a v e s of D elaw are, and
24 ppm and 54 ppm for Ontario le a v e s.

Scott (1944) rep orted that C oncord v in e s

w e r e r e la tiv e ly low in boron, if only 7. 4 ppm B could be found, but that the d e fi­
c ie n c y range w as betw een 10 and 20 ppm a s com pared to the norm al content
range of 15 to 30 ppm.

D eficien t le a v e s exhibited y ello w in g betw een the v e in s

and at the m argin s; no n e c r o s is or p rem a tu re d efoliation o ccu rred , but the
le a f su rfa ce w as abnorm ally rough w ith r a ise d a r e a s betw een v e in s.
in tern od es and tw isted flow er c lu s te r s w ith little fruit s e t w e re noted.

Short

16.

W ilhelm (1952), Branas and Bernon (1954, 1956) and GMrtel (1954) d e scr ib e d
the sam e sym p tom s.

D epending on the s e v e r ity of boron d e fic ie n c y and on

the v a riety , the follow in g sym p tom s have been reported:

tip d ie-b ack , root

d ie-b ack , c u rlin g of le a v e s, m is s in g of in tern od es (double diaphragm ) and
co m p le te ly w hite le a v e s with brow nish sp ots w hich can be seen if held a g a in st
the light.

G&rtel (1956) found that R ie slin g le a v e s w ere d eficien t at 14 ppm B

and had n e c r o s is at 10 ppm B with the tra n sitio n zone being betw een 13 and 15
ppm .
In sand cultu re, M eier (1937), Eaton (1944), Scott and Schrader (1947)
and GMrtel (1956) produced e s s e n tia lly the sam e sym ptom s a s found in v in e ­
y a rd s.

Eaton, by u sin g . 04 ppm B in solution, found 38 ppm B in M alaga

le a v e s and 86 ppm in Sultanina le a v e s, w hich w ere fed with the sam e solution.
Scott and Schrader o b serv ed w a te r-so a k ed a r e a s in the a p ica l te n d rils.

They

found 29 to 41 ppm B in ste m s, and 57 to 146 ppm B in le a v e s of the sam e
p lan ts with boron in solution, but only 20 to 25 ppm B in le a v e s w here no boron
w as added to the solution .

G&rtel (1956) found 8 ppm B in R ie slin g le a v e s

when grown without boron, as com pared to 38 ppm B when 1 m g boron p er
lite r w as added.
A skew (1944) rep orted a com bined c a se of p otassiu m and boron d e fi­
c ie n c y from New Z ealand.

The fir s t sym ptom found on the le a v e s w as a lig h tly

ch lo r o tic m ottlin g betw een the m ain v e in s w hich la te r changed to a red d ish -

17.

brown c o lo r .

At that tim e the m a rg in s of the le a f becam e sco r c h e d and

cu rled upw ards and inw ards.

The b erry settin g w as v e ry ir r e g u la r and

brown t is s u e appeared in the fle s h o f the b e r r ie s .

The boron content show ed

a se a so n a l fluctuation in the le a v e s , as w ell a s in the b e r r ie s .
v a lu e s w ere 10. 3 ppm B and .32%

T he lo w e st

in le a v e s , and 4. 4 ppm B and . 65%

K^O in b e r r ie s .
W ilhelm (1952) and G&rtel (1954) d e scr ib e d boron to x icity sym p tom s.
The le a v e s in th e se c a s e s w e re cupped up or down with r o lle d -in m argin s,
the dentation see m e d to be ligh ter brown than the le a f and the m a rg in s w ere
torn due to the ten sion e x er te d by uneven growth.

L ea v es w hich w ere n o r ­

m a lly fiv e -lo b e d developed only into th ree lo b es.
L eaf a n a ly sis by G&rtel show ed the follow ing resu lts:

healthy le a v e s

28. 2 ppm B, w eakly dam aged 310 to 640 ppm B, and stro n g ly dam aged 620 to
852 ppm B.
Copper:

Rough and unhealthy looking bark, short ca n es, s e v e r e ly

r e s tr ic te d internodal growth, sm a ll, pale, slig h tly ch lo ro tic le a v e s, poor
ro o ts, adventitious bud developm ent near the so il su rfa ce, w ere sym ptom s
of copper d e fic ie n c y d e scr ib e d by T eakle et a l. (1943).
Sultana le a v e s contained,

Copper d eficien t

in th is c a se , 2. 1 to 5. 4 ppm Cu, a s com pared

to 7. 5 to 9. 9 ppm Cu in healthy le a v e s, taken in D ecem b er.
Z inc:

"L ittle L eaf'1, "R eisigkrankheit", "court-noue", or " A rrici-

18.

am ento" are n am es in v a r io u s langu ages for zin c d e fic ie n c y (Dufrenoy,
1935).

B ioletti and Bonnet (1917) d e sc r ib e d the follow in g zin c d eficien cy

sym ptom s:

the a ffected le a v e s show ed tendency to c u rl up and bands or

p a tch es of lig h t-c o lo r e d parenchym a tis s u e could be found.

F u rth erm o re,

a gum m y se c r e tio n in the conducting tis s u e , fla tten ca n e s w ith a v era g e
internode length 50% sh o r te r than norm al, dark sp ots on roots, and non­
se ttin g of fruit w ere o b serv ed .

E ss e n tia lly the sam e d e scr ip tio n s w ere

given by Chandler_£tjrL (1933, 1934), Coombe (1949), C lore (1951), andCook (1957).

Coombe, C lore, and Cook m entioned dw arfing of le a v e s at

shoot tip s, in terv ein a l y e llo w ish to g r e y is h -g r e e n co lo r of le a v e s, and
short shoot grow th.

C lore (1951) com pared the le a f appearance of a zin c

d eficien t plant with one injured by 2, 4 -D .

Cook a s so c ia te d v isu a l d eficien cy

sym p tom s on v in ifera type g ra p es with 15 ppm Zn in the leaf.
M olybdenum :

Basal le a v e s show ing m arginal n e c r o s is developing

c o n c en tr ica lly from the le a f m argin and term in a l le a v e s show ing c h lo r o sis
w e r e rep orted a s m olybdenum d eficien cy by Bergman and Kenworthy (1956).
P e tio le s of p lan ts grow n in nutrient culture, w hich show ed th ese sym ptom s
contained . 2 ppm Mo, a s com p ared to 3. 4 ppm Mo in p lan ts trea ted with
m olybdenum .

The n e c r o s is on b asal le a v e s w a s d e scr ib e d to be the e x ­

p r e s s io n of nitrate to x ic ity inasm u ch a s m olybdenum is e s s e n tia l for the
a s s im ila tio n of n itra tes.

19 .

C hlorine:

W hile th ere has been no d irect b en eficia l e ffe ct on ch lo rin e

rep orted , T hom as (1934) rep o rted the fo llo w in g to x ic ity sym ptom s: p rem atu re
opening of the buds, sm a ll le a v e s and fru its, d efoliation and in e x tr em e c a s e s
faulty fruit set, w h ile m a rg in a l and in ter v e in a l n e c r o s is w e re sym ptom s for
s e v e r e to x ic ity .

R avikovitch and Bichner (1937) found brown sp ots a ll o v er the

le a f blades and y ellow tip le a v e s in addition to the ab ove-m en tion ed sym p tom s.
Under P a lestin e con d ition s d e scr ib e d by the a fo re-m en tio n ed , the NaCl c o n ­
tent v a ried from . 04 to 3. 35%.

L ea v e s of d eterio ra ted C h a sse la s had the

h ig h est NaCl content, w h ereas healthy M uscat Hamburg had . 15%, w h ile
n e a r ly dead on es show ed 1. 5% NaCl. Woodham (1956) felt that p e tio le s w ere
b etter for d eterm ination of the ch lo rid e status than le a f b la d es.

In undrained

loam s o ils of A u stra lia , when th ere w ere slig h t to s e v e r e le a f burns p r e se n t
in D ecem b er, the ch lo rid e content w as betw een . 8 and 1. 9%, and in s e v e r e
c a s e s v a lu es of 2. 5% Cl and m o re w ere found in January.

On healthy s ite s ,

no sym p tom s could be found in the p r e se n c e of 1. 7 to 2. 0% Cl.

T h ere w as

a lw ays som e ch lo rid e p r e se n t w here v in e s w ere affected, but without any
v is ib le sym p tom s.

D ille y (1957a) rep o rted that in c r e a sin g le v e ls o f ch lo rid e

and sulfate in nutrient cu ltu re d e p r e sse d grow th in proportion to the ch lo rid e
le v e l.

The ch lorid e le v e l w a s in each c a se d ir e c tly proportion al to the

supply in the solu tion .

In crea sin g le v e ls of ch lorid e in c r e a se d nitrogen,

copper and zin c, but d e c r e a se d m agn esium in the p e tio le.

20.

Sulfur:

D ille y (1957b) and H arding (1957) found sulfu r d e ficien cy

sy m p tom s on C oncord le a v e s in nutrient cu ltu re w ork by d eletin g the sulfate
ion from the nutrient solution .

At fir s t the term in a l le a v e s turned yellow

and brow nish sp ots appeared on the su r fa c e.
turned into pu rp lish c o lo r in both c a s e s .

T h ese brow nish sp ots la te r

D ille y (1957a) used 192 ppm SO^

in the solution and found . 27% S in the p e tio le, when the SO^ concen tration
w as doubled, he found . 37% S and . 75% S when 624 ppm SO^ w ere added.
C hloride v s. Sulfate:

V inet (1935) concluded, a fter a lo n g -term

study of KC1 v e r s u s K^SO^ fe r tiliz e r , that the la tter w as the better so u rce
for p o ta ssiu m .

High ch lorid e d e c r e a se d the e ffe c tiv e n e s s of p otassiu m

u tilization , w hile su lfate did not.

Shaulis (1954) a lso recom m end ed p otassiu m

su lfate in favor of m u riate of potash.

L a rsen et al_. (1957) rep orted on a

fo u r -y e a r field exp erim en t with both f e r tiliz e r s that fie ld s w ith s e v e r e p o ta s ­
sium d eficien cy r e c o v e r e d fa s te r in the fir s t y ear w ith K SO but in the long
2

cr

run no sign ifican t d ifferen ce in y ield (25 and 26 pounds o f g ra p es p er vine)
and p e tio le com p osition (2. 14 and 2. 08% K) w ere found.

EXPERIMENTAL PROCEDURE

The investigations to study the resp onse of Concord grapes (V itis
labrusca) to various le v e ls of esse n tia l nutrient elem en ts w ere conducted
in the greenhouse in three separate experim ents from January to May,
1956 and 1957.
Experim ent I w as a study of the effects of high and low le v e ls
of eleven nutrient elem en ts on the p etiole com position and growth of grape
p lan ts.
Experim ent II w as a study of the influence of in creasin g le v e ls of
calcium and m agnesium with constant le v e ls of potassium in the solutions
upon p etio le com position and growth of grape plants.

The m anganese -

potassium interrelationship was studied in Experim ent III.
Experim ents II and III w ere conducted sim ultaneously.

22.

EXPERIMENT I
E FF E C T S OF VARIOUS LEVELS OF 11 NUTRIENT ELEM ENTS ON
GROWTH, PETIOLE AND STEM COMPOSITION

In o rd er to find the e ffe ct of nitrogen, ph osphorus, p o ta ssiu m , c a l­
cium , m agn esiu m , iron, m an ganese, boron, copper, zinc, and m olybdenum
on the grow th of Concord gra p es, a random ized block exp erim en t u sin g
nutrient so lu tio n s w as e sta b lish e d .

The exp erim en t contained 25 trea tm en ts

w ith th ree r e p lic a tio n s of each treatm ent.
The nutrient solu tion s for the 25 trea tm en ts w ere obtained by u sin g
Hoagland and Arnon (1950) solution as check, and adjusting the solution to
p rovid e low and high le v e ls of 11 nutrient e lem en ts.

F or low le v e ls the

s p e c ific e lem en t w as c o m p letely deleted from the solution for a ll nutrients
e x cep t nitrogen.

Sm all qu an tities of nitrogen w ere added after four w eek s

in ord er to keep the p lan ts a liv e .

F o r high le v e ls , the sp e c ific elem en t w as

in c r e a s e d in am ounts equal to five tim e s th eir content in Hoagland solution.
High le v e ls of p otassiu m and calciu m w ere studied with two s o lu tio n s -one u sin g sulfate, and the oth er u sin g ch lo rid e a s the anion.
Stock solu tion s w e r e p rep ared by d isso lv in g C. P. grade c h e m ic a ls
in d is tille d w ater, and nutrient solu tio n s w ere m ade up as needed by adding
the n e c e s s a r y stock so lu tion s to ca rb o y s containing 18 lit e r s o f d istille d
w ater (Appendix T a b les 1, 2, 3 and 4).

23.

O n e -y e a r -o ld rooted C oncord grape cu ttin gs w ere dug from a n u rsery
bed n ear Paw Paw, M ichigan, on N ovem ber 12, 1955 and p la ced in cold s t o r ­
age at 40° F until January 28, 1956, w hich provided su fficien t cold to break
dorm ancy, a cco rd in g to M agoon and D ix (1943).
On January 28, 75 p lan ts w ere s e le c te d for the ex p erim en t.
p lan ts w ere u n iform ly pruned to two buds and w eighed.

The

Due to the v a r ia ­

b ility in w eight, it w as n e c e s s a r y to m ake th ree different w eight groups,
having a v era g e w eigh ts of 28, 33 and 41 g r a m s.

The 25 plants in each w eight

group fe ll w ithin one standard deviation, each group com posed a rep lica tio n .
A dditional p lan ts r e p r e se n ta tiv e of each group w ere s e le c te d and u sed to
e sta b lish the o rig in a l dry w eight in order to ca lcu la te la ter the total dry
w eight accu m ulation.
The rooted cuttings w ere planted in 1 2 -inch c la y pots, w hich had
been painted w ith asphaltum paint to prevent contact of ro o ts with the pot
su rfa ce, and red uce evaporation through the s id e s of the p ots.

Watch g la s s e s ,

4 in ch es in d iam eter, w ere p laced in the bottom o f the pots to in su re proper
drainage.

C oarse grade quartz sand w as u sed a s growth m edium .

One green h ou se bench w as u sed p er rep lica tio n .
a ssig n e d at random w ithin each rep lica tio n .

T rea tm en ts w ere

The plants w ere h ea v ily w atered

with d is tille d w ater during the fir s t w eek s and feeding began in the third w eek.
Until the m iddle of M arch, the p lan ts w ere fed th ree tim e s each w eek, one

quart of nutrient solution per pot.
ev ery other day.

T hereafter they w ere fed for one month

From April 1 until term ination of the experim ent, w ater­

ing ev ery day was n ec essa r y .

The plants w ere given nutrient solution two

days in su c cessio n , and d istilled w ater the third day.
When the experim ent w as term inated, the plants w ere harvested by
rep lication s on May 18, 19 and 20.

Total linear growth and dry weight a c ­

cum ulation for p etio les, lea f blades, stem s and roots w ere estab lish ed for
each plant.

Any plant part below the point of pruning at the beginning of

the experim ent w as designated as roots, and the parts above this point, e x ­
cluding p e tio le s and leaf blades, as stem s.

The fibrous root p a rticles were

obtained by p a ssin g the roots through a fine siev e (U. S. 45, 350 m icrons
opening) after drying.
P etio les and stem s from each plant w ere analyzed in the laboratories
of the Department of A gricultural C hem istry.

Nitrogen was determ ined by

the standard Kjeldahl method, and potassium by flam e photom eter.

Phos­

p h o r u s/c a lc iu m , m agnesium , iron, m anganese, boron, copper, and zinc
w ere determ ined spectrographically.
fied method of Samson (1953).

Chloride was determ ined by a m odi­

25.

RESULTS
DEFICIENCY AND TOXICITY SYMPTOMS OF STEMS AND LEAVES

Under norm al conditions a plant seld om show s v isu a l d e fic ie n c y or to x i­
c ity sym p tom s on le a v e s , excep t under ex trem e s t r e s s for one or m o re e le m e n ts.
In th is exp erim en t, d e fic ie n c y sym ptom s on le a v e s w ere o b se rv e d for the low
le v e l of each of the 11 e le m e n ts.

On the oth er hand, only four of the 13 high

le v e l trea tm en ts exh ib ited le a f sym p tom s.

T h ese sym ptom s m ight have been

due to to x ic ity of the individual elem en t, or w ere c h a r a c te r istic of d eficien cy
sym p tom s of other e le m e n ts, induced by the high le v e l of the elem en t being
app lied in e x c e s s .
G eneral O b serv a tio n s:

A few of the plants, h e a v ie st in in itia l w eight,

produced flo w e r s sh o rtly after in itiation o f growth.

T h e se flo w e r s w ere r e ­

m oved in ord er to p reven t any drain of nutrients a sso c ia te d w ith fruit fo r m a ­
tion.
The plan ts of the lig h te st w eight group began to grow m o re rapidly than
p lan ts of the oth er two groups.

H ow ever, a fter fiv e w eek s a ll plan ts w ere r e la ­

tiv e ly ev en in growth, and at the term in ation of the ex p erim en t the plan ts of
the h e a v ie st w eight group had m ade m o st lin ea r growth, if not too s e v e r e ly
a ffected by the v a rio u s tr e a tm e n ts.
V isu al d e fic ie n c y or to x ic ity sym p tom s on le a v e s appeared v ery slow ly,
ex cep t for the low n itrogen p lan ts on w hich s e v e r e sym ptom s w e re p r e se n t

26.

a fter 4 w eek s of grow th.

The oth er trea tm en ts produced le a f sym p tom s in

the fo llo w in g ch ron ological order:

5 N and 5 P a fter 6 w eek s; - K and 5 K (SO^)

a fter 9 w eek s; 5 K (Cl) and - F e after 10 w eek s; follow ed by - Mg, - Mn, - B,
and - Cu one w eek la te r .

- P, - Ca and - Zn had v isu a l sym p tom s a fter 12

w eek s of growth, and - Mo a fter 14 w eek s.

A ll oth er trea tm en ts did not show

v isu a l le a f sym p tom s, but lin ear grow th and dry w eight accum ulation w ere in
so m e c a s e s v e r y s e v e r e ly affected .
N itrogen:

The plan ts supplied w ith nutrient solution d eficien t in n itr o ­

gen show ed a v is ib le reduction in rate of lin ea r grow th after only 3 w eek s.
L e a v e s produced during th is tim e showed a healthy g reen co lo r, but began to
put on a ligh t g reen c a st w hich sh o rtly turned o liv e and fin ally y ello w ish
g reen .

At th is point, term in a l grow th stopped co m p letely .

Dark brown p in ­

head lik e sp ots app eared on the upper su rfa ce o f the basal le a v e s (F igure 1,
top).

T h e se sp ots extended through the le a f and w ere v is ib le on the low er

e p id e r m is.

At fir s t it w as fe lt that th ese spots m ight be due to in se c t injury

o r pathogens.

Thorough exam ination by plant p ath ologist and en tom ologist,

how ever, ruled out e ith e r of th ese p r o b a b ilitie s.
m in us nitrogen p lan ts exh ib ited th is sym ptom .

On the other hand, only the

At th is tim e 42 ppm of nitrogen

w as b ein g added to the nutrient solution in o rd er to prevent co m p lete sta r v a ­
tion.

The brown sp ots d isap p eared w ithin a short tim e and the le a f co lo r r e ­

turned to a light g reen .

T erm in al grow th began anew and new le a v e s,

not

F ig u r e 1.

Top

G rape le a f from the b a s e o f a n itr o g e n
d e fic ie n t p lan t, a fte r th r e e w e e k s o f
grow th . T he p in -h e a d lik e d ark brow n
sp o ts w e r e o b s e r v e d on the a d a x ia l and
a b a x ia l s u r fa c e s o f the le a v e s .

Bottom

G rape p la n t su p p lie d w ith h ig h le v e l s
(1050 ppm ) o f n itr o g e n in th e n u trien t
so lu tio n . T he plant g r ew p r o fu s e ly and
p ro d u ced la r g e , v e r y d ark g r e e n le a v e s .
T he te r m in a ls tu rn ed c h lo r o tic a fte r 10
w e e k s o f grow th.

27.

28.

quite a s la r g e a s th ose of the ch eck plants, developed.
le a v e s dropped off la te r .

Som e of the basal

M ost of the le a v e s on the lo w er o n e-th ird of the

sh o o ts w e re lik e heavy paper, v e ry stiff in tex tu re.
fo rm ed during the rem ain in g grow ing sea so n .

No la te r a l shoots w ere

Final ch em ica l a n a ly sis of

p e tio le s show ed . 51% nitrogen, com pared to 1. 06% N in p e tio le s of the check
p lan ts.
Plants supplied w ith high le v e ls of nitrogen grew p r o fu se ly and p r o ­
duced la r g e, v e ry dark g reen le a v e s .

A fter 10 w eek s, young term in al le a v e s

app eared ch lo r o tic (F igu re 1, bottom ), w ith each new le a f from then on hav­
ing the sam e ap p earan ce.

Shortly b efore term in ation of the exp erim en t, the

youn gest le a v e s w e re v e r y wavy in addition to being ch lo ro tic.

A sym ptom

s im ila r to the one d e scr ib e d appeared at the sam e tim e on p otassiu m d eficient
p lan ts.

P etio le a n a ly sis show ed 5. 19% nitrogen, and 1. 57% p o ta ssiu m ,

w hich w as the lo w est p o ta ssiu m content of any treatm en t ex cep t for the low
p o ta ssiu m treatm en t.

H ow ever, the o c cu rr e n c e of p otassiu m d eficien cy

sym p tom s on term in a l le a v e s see m e d q u estion able.

The p o s s ib ility of zin c

d e fic ie n c y e x iste d and w ill be d is c u s se d in connection with p o ta ssiu m .
P h osp horus:

Plants grown without phosphorus show ed no v isu a l s y m ­

ptom s on the le a v e s fo r e lev e n w eek s.

The lin ea r growth, how ever, w as

rath er slow and r e s tr ic te d , and c e a s e d co m p letely in the late stage of the
ex p erim en t.

The le a v e s, in g en era l, w ere dark green in co lo r and v ery

29.

h eavy or le a th e r y in tex tu re.

A ctually, the fir s t v isu a l sym ptom on the le a f

w a s the red c o lo ra tio n of v e in s on he abaxial sid e of the b asal le a v e s (F ig u re
2 , bottom left), w hich w as follow ed by the d isap p earan ce of the g reen pigm ent
from the le a f su rfa ce, beginning at the m argin and p r o g r e s s in g cen trip eta lly ,
a s shown in F ig u re 2 top and bottom right).
sta y e d on the plant and did not drop off.

The a ffected le a v e s, how ever,

L ater p e tio le a n a ly sis show ed . 10%

p h osp horus content in p e tio le s of th ese plan ts, as com pared to . 37% in
p e tio le s of the c h eck s.
The p lan ts supplied w ith high le v e ls of phosphorus developed norm ally
in e v e r y r e sp e c t, and v e ry little d ifferen ce could be o b serv ed betw een th ese
p lan ts and the check p lan ts.

A fter 6 w eeks, how ever, term in a l le a v e s on

la te r a l sh oots w ere slig h tly ch lo r o tic .

T his sym ptom never developed into

a s p e c ific pattern and a d ia g n o sis based on it would not have been r e lia b le .
P etio le a n a ly sis did not r ev e a l, for certain , the ca u se of the c h lo r o sis.
Sin ce 1. 41% ph osphorus and . 64% calciu m

w ere found in the p e tio le, a sligh t

c a se of calciu m d e fic ie n c y could be su sp ected .
P otassiu m :

A fter 9 w eek s of slow growth, the b asal le a v e s on one

plant grown without p o ta ssiu m began to show slig h t yellow in terv ein a l c h lo r o sis
that extended to the le a f m argin.
n e c r o tic .

T he m argin o f th ese le a v e s la ter turned

The le a v e s on the oth er two p lan ts, how ever, did not show th is

sym ptom but a fter 11 w eek s had so m e p u rp lish b lotch es (F igu re 3, bottom

F ig u r e 2.

Top

B asal p o r tio n o f a g r a p e p la n t su p ­
p lie d w ith p h o sp h o r u s d e fic ie n t nutrient
so lu tio n a s o b s e r v e d a ft e r 11 w e ek s of
grow th.

Bottom le ft

A b a x ia l s id e o f a b a s a l le a f sh ow in g
red c o lo r a tio n o f v e in s , in d ic a tiv e of
p h o sp h o ru s d e fic ie n c y .

Bottom righ t

A d a x ia l s id e o f the s a m e b a s a l le a f.

31 .

right) on the adaxial surface of the basal lea v es.

Tip lea v es, in both c a se s,

began to exhibit yellow brownish interveinal ch lo ro sis after 11 weeks, with
v ery wavy m argins (Figure 3, top, and bottom left).
nearly at a standstill.

Stem elongation was

All plants had very short internodes and the p etioles

w ere strikingly short, as found in vineyards having sev ere potassium defi­
ciency.

Petiole an a ly sis showed a very low . 12%, potassium content, which

com bined with 25 ppm zinc, w as the low est content for either elem ent in any
treatm ent.

The ch lo ro sis of the term inal leaves could be suspected to be an

ex p ressio n of zinc deficiency.
Two treatm ents receiv ed high le v e ls of potassium , one contained
potassium sulfate and the other contained potassium chloride in the solution.
Both treatm ents produced the sam e visual leaf sym ptom s.

After 9 weeks of

growth, the potassium sulfate treatm ent began to show brown irregular
blotches in the interveinal field s of the basal lea v es (Figure 4, bottom left).
At fir st, because of very high sulfate content of the nutrient solution (1342
ppm SO^), sulfate toxicity was expected, but then one week later the sam e
sym ptom s appeared on the potassium chloride treatm ents, except the basal
lea v es w ere not fir st affected but rather those of the fourth and fifth in ternodes (Figure 4, top right).

The blotches enlarged quite rapidly, began to

join each other, and p ro g ressed centrifugally.

The veins of th ese leaves

w ere green at all tim es and a green m argin was always p resen t (Figure 4,

F ig u r e 3.

Top

G rap e p lan t su p p lie d w ith p o ta s s iu m
d e fic ie n t n u trien t so lu tio n , a s o b s e r v e d
a fte r 9 w e e k s o f grow th: c h lo r o tic le a v e s
at th e te r m in a ls and p u r p lis h b lo tch ed
le a v e s at the b a s e .

Bottom righ t A d a x ia l s u r fa c e o f a p o ta s s iu m d e fic ie n t
b a sa l le a f w ith p u r p lis h b lo tc h e s .
Bottom le ft

L ea f fro m te r m in a l p o r tio n o f a p o ta ssiu m
d e fic ie n t p la n t. P e tio le a n a ly s is su g g e ste d
z in c d e fic ie n c y .

F ig u r e 4.

Top le ft

G rap e plan t su p p lie d w ith h ig h l e v e l s (1171
ppm ) o f p o ta s s iu m s u lfa te , a fte r 9 w e e k s of
grow th .

Top rig h t

G ra p e plan t su p p lie d w ith high l e v e l s (1171
ppm ) o f p o ta s s iu m c h lo r id e , a fte r 10 w e e k s
o f grow th .
N ote th e id e n tic a l sy m p to m s on both p la n ts.

Bottom le ft

F i r s t v is u a l sy m p to m s o f n u tr itio n a l d iso r d e r
a s o b s e r v e d on p la n ts su p p lie d w ith h ig h le v e ls
o f p o ta ss iu m w ith su lfa te o r c h lo r id e . T h is
s a m e sy m p to m a ls o a p p e a r e d on le a v e s from
th e m id - s e c t io n o f p la n ts su p p lie d w ith m a g ­
n e siu m d e fic ie n t n u tr ien t so lu tio n ( s e e F ig u r e
5, top).

Bottom rig h t

T he sa m e le a f a s on bottom le ft, a few days
la te r . T he v e in s w e r e g r e e n at a ll t im e s and
a g r e e n m a r g in w a s a lw a y s p r e s e n t . T he
sy m p to m w a s id e n tifie d by p e t io le a n a ly s is
a s m a g n esiu m d e fic ie n c y .

33 .

34 .

bottom right).

The brown tissu e slow ly dried out com pletely and the affected

le a v e s fe ll off (Figure 4, top).

On the potassium chloride treatm ent, this

symptom spread both acrop etally and basipetally.

At the sam e tim e, s im i­

lar sym ptom s appeared on m agnesium deficient plants, and therefore both
sulfate and chloride toxicity w ere ruled out in favor of m agnesium deficiency.
T his w as later v erified by the p etiole an alysis, inasm uch as both high level
potassium treatm ents had a petiole content of . 14% m agnesium and a very
high potassium content of 9. 86% for the potassium sulfate treatm ent and
8. 88% for the potassium chloride treatm ent.
M agnesium :

Plants grown without m agnesium developed norm ally

like the check plants and no difference could be seen until the 11th week
when the basal lea v es began to turn yellow in the interveinal spaces of the
adaxial side (Figure 5, bottom).
p ro g r ess to the lea f edge.

The yellow pigment, however, did not

Figure 5 (top) shows a m agnesium deficient plant

with two different m agnesium deficiency sym ptom s, one being the yellow
interveinal coloring on basal leaves and the other one being like the d e s ­
cription of the high potassium sym ptom s (Figure 4, bottom), which developed
sim ultaneously.

M agnesium deficient plants grown with m agnesium deficient

solution had . 06% m agnesium in the p etiole.
Under high le v e ls of m agnesium , no visu al leaf sym ptom s could be
observed.

The linear growth seem ed, however, very much depressed.

F ig u r e 5.

Top

Bottom

G rap e p lan t su p p lie d w ith m a g n e siu m d e fic ie n t
n u tr ien t so lu tio n , a fte r 11 w e e k s o f grow th.
T w o d is tin c t le a f sy m p to m s a p p e a r e d s im u l­
ta n e o u sly , o n e a t th e b a s e and the o th e r at the
m id - s e c t io n o f th e p la n t. T h e sy m p to m on the
m id - s e c t io n le a f w a s th e sa m e a s o b s e r v e d on
p la n ts su p p lie d w ith h ig h l e v e l s o f p o ta ssiu m
(s e e F ig u r e 4, b o tto m ).
M a g n esiu m d e fic ie n t le a f w ith y e llo w in t e r ­
v e in a l c h lo r o s is fro m the b a s a l p a r t o f the
p lan t.

35 .

36.

C alcium :

The appearance of calciu m d eficien cy sym ptom s on p lan ts

grow n without calciu m w as actu a lly ex p ected to occu r m uch soon er.

The

plan ts se e m e d to do w e ll up to the 11th w eek, at w hich tim e the d istilla tio n
apparatus broke down.

D uring rep air, a heavy d ep osit of calcium w as r e ­

m oved from its inner w a lls.

Within one w eek, a d ifferent appearance of the

youn gest le a v e s at the term in a l could be o b serv ed .

W hereas norm al grape

le a v e s have an equally lobed base, the new le a v e s had a co rd a te-o b lo n g b ase.
A lso , the n orm ally shallow toothed m a rg in s w ere in th is c a se prom in en tly
toothed and the w hole le a f had a rather oblong form .

Soon th ese young

le a v e s turned yellow and blackening of v ein s on the abaxial sid e of the lea f
could be ob served , a s shown in F igure 6 (bottom left).

Sim ultaneously,

la rg e w a te r -so a k e d a r e a s in the apical p ortion s of the te n d rils appeared
w hich led to the dieback of the tend ril at fir s t, follow ed by dieback of the
w hole term in a l p ortion of the shoot to the next internode (F igure 6, top right).
Growth of the a x illa r y buds (F igu re 6, bottom right) at th ese im m ed iate in te r ­
nodes w as brought about by the dieback.

By now, old er le a v e s exhibited in ter-

vein a l c h lo r o sis com bined w ith cu rlin g under the lea f blade, givin g it a beak­
lik e appearance, and a ll te r m in a ls, r e g a r d le s s of th eir location, began to
break down (F ig u r e 6, top right).

U ltim ately, the o ld est portion of the ste m s

b ecam e p itted in app earance, looking sim ila r to s e v e r e hail dam age cau sed by
v e r y fine hail sto n es (F igu re 6, bottom right).
p e tio le s w as . 26%.

The calcium content of th ese

F ig u r e 6

Top le ft

G rape p la n t su p p lie d w ith c a lc iu m d e fic ie n t
n u trien t s o lu tio n in th e b reak d ow n s ta g e .
T he plan t w a s v e r y s e v e r e ly d am aged , w ith
a ll te r m in a l p o r tio n s sh o w in g d ie -b a c k .

Top rig h t

T e r m in a l p o r tio n o f a c a lc iu m d e fic ie n t
g r a p e . N ote the b e a k -lik e a p p e a r a n c e of
th e le a f, th e w ith e r in g te n d r ils , and the
d e v e lo p m e n t o f a x illa r y b u d s.

Bottom le ft

B lack en in g o f the v e in s on th e a b a x ia l sid e
and y e llo w in g o f the le a f, an e a r ly in d ic a tio n
o f c a lc iu m d e fic ie n c y .

Bottom rig h t

G row th o f an a x illa r y bud brough t about by
d ie -b a c k o f te r m in a l p o r tio n s . N ote the
p ittin g o f the ste m and d e v e lo p in g o f d o u b le bud.

37 .

38.

High le v e ls of calciu m w ere te s te d w ith two different nutrient s o lu ­
tio n s, one having su lfate and the other having ch lo rid e a s the anion c a r r ie r
of c a lciu m .

On n eith er treatm en t w ere v isu a l le a f d e ficien cy or to x icity

sym p tom s o b serv ed .

A ctually, s e v e r e ch lorid e injury w as exp ected w here

calciu m ch lorid e w as u sed b ecau se of the p r e se n c e of 1418 p p m /lite r of
ch lo rid e, but only a s e v e r e d e p r e ssio n in lin ea r growth occu rred .

The

calciu m content of p e tio le s from plants treated with calcium sulfate w as . 96%
Ca com p ared to 2. 55% ca in p e tio le s of plan ts trea ted with calcium ch lorid e.
Iron:

Plants grow n w ith low le v e ls of iron produced slig h tly ch lo ro tic

le a v e s at the ap ical p ortion s of the m ain ste m s and young shoots a fter ten
w eek s of grow th.

The sym ptom w as, how ever, n ever pronounced enough to

be c o n sid e re d of im p ortan ce.

The sam e held true for plants trea ted with high

le v e ls of iron, in w hich c a se only lin ea r growth w as d ep ressed .
M anganese:

A fter 11 w eek s of growth, plants grown without m anganese

in the nutrient solution began to show light g reen co lo r in the in tervein al field s
of young ap ical le a v e s of both m ain and la tera l sh oots.

The v e in s th e m se lv e s,

how ever, n ever lo st th eir norm al g reen co lo r (F igure 7, bottom left).

P e tio les

of m an gan ese d eficien t p lan ts had only an 18 ppm m an ganese content com pared
to 63 ppm in the check .
The high le v e l m an gan ese plan ts grew lik e norm al plan ts without any
sig n s of varia tio n .

The p e tio le a n a ly sis in th is c a se rev e a le d 363 ppm Mn,

F ig u r e 7

Top le ft

T e r m in a l p o r tio n o f a g ra p e p la n t su p p lied
w ith b oron d e fic ie n t n u tr ien t so lu tio n , a fte r
11 w e e k s o f g ro w th . T he in te r n o d e s w e r e
v e r y sh o rt and th e te r m in a l a p p e a r e d stunted.
T he te n d r ils sh o w ed w a te r - s o a k e d s p o ts .

Top rig h t

T e n d r il o f the sa m e p la n t sh o w in g w a te r so a k ed sp o ts, th e b e g in n in g o f b reak d ow n
and an e a r ly in d ic a tio n o f s e v e r e boron
d e fic ie n c y .

Bottom le ft

Y oung te r m in a l o f a g r a p e p la n t su p p lie d w ith
m a n g a n e se d e fic ie n t n u tr ien t so lu tio n , a fte r
11 w e e k s o f grow th .

Bottom righ t

O ld er le a f fro m b oron d e fic ie n t p lan t a s o b ­
s e r v e d fro m th e a b a x ia l s id e a g a in s t su n ­
lig h t.

40.

w hich is n ea rly equal to a norm al fie ld sam ple content and it is, th erefore,
e a s ily understandable why no sym ptom s occu rred .
Boron:

Plants grown without boron m ade a stunted appearance after

11 w eek s of grow th (F igu re 7, top left).
and v e r y s tiff in tex tu re.

The term in a l le a v e s w ere yellow

W hile w ith calcium d eficien cy th ese le a v e s turned

from green to yellow , boron d eficien t le a v e s d iffered by being yellow from
the tim e they unfolded.

No blackening of the v e in s o ccu rred at any tim e

w ith boron d eficien cy , but a s it w as with the c a se of calcium d eficien cy, w a tersoaked sp ots appeared in the ten d rils, as shown in F igu re 7 (top right).

The

old er yellow le a v e s se e m e d to be rath er thin, and if held again st strong sun­
light, the v e in s appeared red d ish and the im p re ssio n a r o se that light could
be see n through the le a f (F igure 7 , bottom right).

L ater on, ir re g u la r

y ello w -b ro w n ish a r e a s form ed on the su rface of th ese le a v e s.

The p e tio le s

of th ese le a v e s contained 14 ppm boron.
No v isu a l sym p tom s from the high le v el of boron w ere o b serv ed on
any of the plants r e c e iv in g that treatm ent.
Copper:

Young unfolding le a v e s of C oncord grap es have a p u rp lish -

red d ish tinge, w hich d isa p p ea rs a s soon as the le a v e s produce chlorophyll.
Plants grown without copper behaved the sa m e for about 11 w eek s, at which
tim e the young le a v e s retain ed th is red dish tinge, rather than lo sin g it.

T h ese

le a v e s w ere v e ry fine in textu re and the la te r a l shoots that developed w ere

41.

spindly (F ig u re 8, top left).

A s the le a v e s becam e o ld er the red d ish tinge

changed into a golden co lo r w ith the v e in s turning g reen co lo r, a s shown in
F ig u re 8 (bottom left).
No v isu a l e x p r e s s io n o f the high le v e l of copper w as o b serv ed on plan ts
r e c e iv in g that treatm en t.
Z inc:

L eaf sym ptom s of zin c d eficien cy becam e v is ib le after 12 w eek s

of grow th without a supply of zin c.

At fir st, the young term in a l le a v e s showed

a y e llo w -b r o w n ish in tervein al c h lo r o s is .

The internodal growth slow ed down

con sid erab ly, and the en tire e x p r e ssio n of the term in al le a v e s changed.
m al C oncord le a v e s have an equally lobed b ase and pinnate v e in s.
how ever, the b ase changed to acute and the v ein s w ere p alm ate.

N or­

In th is ca se ,
A lso, the n o r ­

m al sh allow -tooth ed m argin s becam e prom in en tly toothed and v ery wavy.

Both

the term in a l and the la te r a l lobe apex grew acu m in ately in stead of acute and the
la te r a l sinus, w hich is n orm ally o b scu re and so m e tim e s notched, becam e deeply
notched, as shown in F ig u re 8 (top and bottom right).
The high le v e l of zin c produced no v isu a l sym ptom s on the plants.
Molybdenum:

Plants grown without molybdenum developed norm ally and

only a fter 14 w eek s som e slig h tly ch lorotic term in al le a v e s appeared.
s is w as e x p r e s s e d by yello w -b ro w n ish a r ea s on the lea f su rfa ce.
produced n e c ro tic m argin s on a few b asal le a v e s .

The c h lo r o ­

Only one plant

Both sym ptom s w ere not a s d is ­

tin ctly e x p r e s s e d a s d e scr ib e d by Bergman and Kenworthy (1956).
The high le v e l of m olybdenum produced v ery healthy looking plants with
la r g e le a v e s and no ill e ffe c ts w ere ob served .

F ig u r e 8.

Top le ft

M id -s e c t io n o f a g r a p e p la n t su p p lie d w ith
co p p er d e fic ie n t n u trien t so lu tio n , a fte r 11
w e e k s o f grow th. N ote the sp in d ly appearance
o f sh o o ts .

Top righ t

T e r m in a l p o r tio n o f g ra p e su p p lie d w ith z in c
d e fic ie n t n u trien t s o lu tio n .

Bottom le ft

G rap e le a f sh o w in g c o p p e r d e fic ie n c y s y m p ­
to m s. T he le a v e s w e r e g o ld e n c o lo r e d , but
v e in s r e m a in e d g r e e n .

Bottom righ t

T e r m in a l p o r tio n o f a z in c d e fic ie n t g ra p e
p la n t w ith w avy le a v e s sh o w in g y e llo w b ro w n ish in te r v e in a l c h lo r o s is , deep notched
s in u s e s , and p r o m in en t a p e x lo b e s .

43.

VISUAL SYMPTOMS ON ROOTS

The g en era l appearance of root s y s te m s v a ried g rea tly betw een tr e a t­
m en ts.

It w as fe lt that the ch eck treatm en t produced ro o ts and a root sy stem

d e sir a b le for plan ts of th is age and, th erefo re, ser v e d a s a b a sis for co m p a r­
iso n .

The effect of the individual trea tm en ts on the ro o ts is shown in Table 1.
G en eral A ppearance of Root S y ste m :

and g en eral appearance w as u sed a s index.

Amount of ro o ts, co lo r, textu re
It w as p o s s ib le that trea tm en ts

having sm a ll r o o ts but looking healthy w ere given a "good".
14 w e re judged a s "good".

Of 25 treatm en ts,

Two trea tm en ts, - Ca and 5 F e, had v ery poor, and

two trea tm en ts 5 P and 5 Mo, had "excellent" root s y ste m s .

5 N, - P and - F e

produced rath er poor root s y ste m s and - N, 5 B and 5 Cu produced v ery good
s y s te m s .

Two trea tm en ts, - Ca and 5 Fe, had v e ry poor appearing root s y ste m s

w ith b r ittle ro o ts interw oven with many dead o n es.
Root Color:
of the r o o ts.

The gen era l appearance w as c lo s e ly rela ted with the co lo r

W hile m o st roots w ere brown, it seem ed that with p r o g r e ssiv e

higher ratin gs of the w hole root sy ste m , the co lo r of the roots becam e ligh ter.
The two "excellent" root s y s te m s of 5 P and 5 Mo, as w ell a s - Mo and 5 B,
which w ere judged good and v e ry good r e sp e c tiv e ly , w ere light brown.
the other hand, - Ca and 5 F e trea tm en ts produced black ro o ts.
of - P and 5 Ca (Cl) w ere of red d ish co lo r.

On

The roots

44.

TABLE 1
VISUAL E F F E C T OF INDIVIDUAL TREATM ENTS ON ROOTS

T reatm en t

G en eral A p p ear­
ance of Root System

C olor of
R oots

R em ark s

Check

Good

Brown

B asis for com p arison
(Standard)

- N
5 N

V ery good
Poor

Brown
Brown

Many fib rou s ro o ts
Few ro o ts, m any dead

- P
5?

Poor
E x ce lle n t

Sm all ro o ts
R eddish
Light brown

- K
5 K (SO )
5 K (Cl)

Poor
Good
Good

Brown
Brown
Brown

V ery few ro o ts
Springy ro o ts, som e dead
Spring ro o ts, m any fibrous

- Ca
5 C a(S04 )

V ery poor
Good

Black
Brown

5 Ca (Cl)

Good

R eddish

V ery b rittle ro o ts, so m e dead
Compact sy stem with many
fib rou s roots
Not too m any roots but v e ry
thin

- Mg
5 Mg

Good
Good

Brown
Brown

Sm all root sy ste m
V ery la rg e sy stem of w iry
roots

- Fe
5 Fe

Poor
V ery poor

Black
Brown

Som e ro o ts w ith w hite tip s
V ery b r ittle and m any dead
ro o ts

- Mn
5 Mn

Good
Good

Brown
Brown

■- B
5 B

Good
V ery good

Brown
Light brown Sm all root sy stem

- Cu
5 Cu

Good
V ery good

Brown
Brown

- Zn
5 Zn

Good
Good

Brown
Brown

- Mo
5 Mo

Good
E x ce lle n t

Light brown
Light brown V ery la r g e com pact sy stem

Som e ro o ts w ith w hite tip s

B rittle ro o ts
Compact sy stem w ith m any
fib rou s ro o ts

45.

T extu re of R o o ts:

On a few trea tm en ts so m e deviation in the te x ­

tu re of ro o ts from the norm al appearance w as o b serv ed .

- Ca, 5 F e and

- Zn ro o ts se e m e d to be b rittle, w hile roots of the two high p otassiu m
tr e a tm e n ts w e re rath er springy, and the 5 Mg roots w ere w iry.

5K (S0 4) |
5K(CI)i
-Co
5C o (SO4 )
5Co(CI)

46 .

GROWTH

Data p erta in in g to total lin ea r length growth, as w ell a s total dry
w eight accu m ulation of ro o ts, ste m s, p e tio le s , and lea f blades a re given
in Appendix T able 5.

Dry w eight accu m ulation of ro o ts, ste m s, p e tio le s,

and le a f b lad es, co n v erted into p er cent o f total dry w eight accum ulation
and sh o o t/r o o t r a tio s a r e record ed in Appendix T able 6.

T otal L in ear Growth (s e e figu re opposite)
The ch eck treatm en t produced the g r e a te st amount o f lin ea r growth
(625 cm ) and a s m ay have been expected , the - N treatm ent produced le a st
lin ea r grow th (109 cm ).
When the ch eck w as com pared s ta tis tic a lly with the individual tr e a t­
m en ts, 5 N, 5 Ca(SO^), 5 Fe, - B and - Cu w ere sig n ifica n tly sm a lle r at the
5% le v e l and - N, - P, - K, 5 K (Cl), - Ca, 5 Ca (Cl) and 5 M g w ere sm a lle r
at the 1% le v e l.
T h ere w as a g en era l tendency for the high le v e l of the nutrients to
r e su lt in le s s reduction of grow th than the low le v e l.
found for m agn esium , iron, and m an gan ese.

T h is tendency w as not

H owever, th ere w ere but few

in sta n ce s w h ere th ere w as a sig n ifica n t d ifferen ce betw een the low and high
le v e l of a nutrient.
grow th than 5 P.

The - P treatm en t produced sig n ifica n tly le s s lin ea r

The high le v e l of m agn esium produced sig n ifica n tly le s s

Check

5K (S04)
5K(C I)

5Ca (S04)

5 C q (CI)

—Mn

-M o

60

80

100

TOTAL DRY WEIGHT ACCUMULATION — (GRAMS)
LSD

5%
I %

—

—

■

Sig. different _ * a t 5 %
than check
# # at I %

120

47.

lin e a r grow th than - Mg.

Both - K and 5 K (Cl) produced le s s growth than

5 K (S 0 4).

T otal D ry W eight A ccum ulation (s e e figu re opposite)
The 25 trea tm en ts produced a la rg e v a riation in total dry w eight a c cu m ­
ulation.

The - K treatm en t resu lted in the le a st production of dry m atter

(35. 45 g).

The high le v e l of Mo produced the g r e a te st amount of dry m atter

(117. 75 g), w hile the check treatm ent produced 90. 70 g of dry m atter.
When the ch eck w as com pared s ta tis tic a lly with the individual tr e a t­
m en ts, only 5 Mo w as sig n ifica n tly higher at the 5% le v e l.

The - P and - B

trea tm en ts w ere low er than the check at the sam e le v e l of sig n ifica n ce.

A 1%

le v e l of sig n ifica n t d ifferen ce w as found for - N, 5 N, 5 P, - K, 5 K (Cl), - Ca,
5 Ca (Cl), and 5 Mg.

The 5 K (S 0 4), - Mg, - Fe, 5 Fe, - Mn, 5 Mn, 5 B, - Cu,

5 Cu, - Zn, 5 Zn, and - Mo treatm en ts did not differ sig n ifica n tly from the
check.

H ow ever, a ll trea tm en ts excep t - Fe, - Cu, 5 Cu, and 5 Zn w ere s i g ­

n ifican tly below 5 Mo in the production of dry m atter.
High and low le v e ls of m agn esium , iron, and molybdenum w ere s ig n i­
fican tly d ifferen t from ea ch oth er.

T h ere w as a significan t d ifferen ce betw een

the 5 K treatm en t u sin g su lfate and the 5 K treatm ent u sin g chloride, a lso
betw een the 5 K treatm en t u sin g su lfa te and the - K treatm en t.

No s ig n ifi­

cant d ifferen ce w as found betw een the - K treatm en t and the 5 K treatm ent
u sin g ch lo rid e.

ROOTS

STEM

20

30
40
O
10
20
2.5 5 0
10
DRY WEIGHT ACCUMULATION — (GRAMS)

PETIOLES

LEAF BLADES

Check

5K(S0«)
5K(CI)P
-C
5Ca(S04
5Ca(CI)

LSD

5%
Sig. different
than check

*a t 5%
# *a t I %

30

40

48.

D ry W eight A ccu m ulation by R oots, Stem s, P e tio les and L eaf Blades (se e
figu re op p osite)
R o o ts:

The 5 K (Cl) treatm ent produced the le a st root dry w eight

(9. 85 g), with the check being in the m iddle (23. 65 g) and 5 Mo producing
m o st root dry w eight (49. 35 g).
When the ch eck w as com pared sta tis tic a lly w ith individual trea tm en ts,
only - Cu and 5 Mo w ere sig n ifica n tly higher, the fo rm er at the 5% le v el and
the la tter at the 1% le v e l.

None of the trea tm en ts produced sig n ifica n tly le s s

root grow th than the ch eck treatm ent.
C om paring the low and high le v e ls of the sam e elem en t, only the low
le v e l of m olybdenum w as sig n ifica n tly below its high le v e l in root growth.

A

com p arison of root dry w eight for the v a rio u s nutrient elem en t le v e ls showed
that th ere w as a g en eral trend for the m ajor elem en ts to produce le s s root
grow th at both the low and high le v e ls than the check treatm ent.

An opposite

trend w as o b serv ed for the m in or e le m e n ts--b o th low and high le v e ls resu lted
in g r ea ter root grow th than the ch eck treatm en t.

An exception to th is trend

w as found for phosphorus w here th ere w as le s s root growth with each increm en t
in the phosphorus supply.
Stem s:

The - K treatm en t produced le a st stem dry w eight (7 g), - Fe

m o st (28. 75 g), and the ch eck 25. 6 g.

49.

A co m p a riso n of the dry w eight accu m ulation of ste m s for the v a rio u s
le v e ls of a nutrient elem en t show ed a g en era l trend for growth to be reduced
by both low and high le v e ls of each e lem en t.

An exception w as found for

m olybdenum w here each in crem en t in supply resu lted in additional stem growth,
w h erea s each in crem en t o f iron resu lted in additional d e c r e a se in stem growth.
The follow in g trea tm en ts w ere sig n ifica n tly low er (5%) than the check:
- N, - K, 5 K (Cl), 5 Ca (SO4), 5 Ca (Cl), and 5 Mg.

W hereas, the 5 K (Cl)

treatm en t produced sig n ifica n tly le s s stem growth than the check treatm ent,
th ere w as no sign ifican t red uction w ith the 5 K (SO^) treatm en t.
m ent, h ow ever, w as sig n ifica n tly la r g e r than the check.

No tr e a t­

A lso none of the

m in or elem en t trea tm en ts resu lted in sig n ifica n tly le s s stem growth than
w as found for the ch eck treatm en t.

The low le v e l of m agnesium resu lted in

sig n ifica n tly m ore stem growth than the high le v e l of m agn esium .
P etio les:

The - K treatm en t produced le a st p etio le dry w eight (1. 1 g),

- F e m o st (5. 11 g), and the check 4. 3 g.
The sa m e gen eral trend a s o b serv ed for stem dry w eight w as a lso
p r e se n t in p e tio le grow th.

Nine of 11 e lem en ts produced le s s p e tio le growth

at both the low and high le v e ls than the check treatm ent.

With iron, th ere

w as a d e c r e a s e in dry w eight of p e tio le s with in c r e a se in ea ch increm en t in
supply, w h erea s with each in crem en t in the supply of copper th ere w as an
in c r e a s e in p e tio le grow th.

50 .

When the ch eck w as com pared s ta tis tic a lly w ith individual trea tm en ts,
no treatm en t w as sig n ifica n tly high er and only the - N, - K, and 5 Mg tr e a t­
m en ts w e r e lo w er than the check .

The 5 K (SO^) treatm en t resu lted in s ig n i­

fican tly m ore p e tio le grow th than the - K treatm en t.

The 5 Mg treatm ent

produced le s s p e tio le grow th than the - Mg treatm en t.
L eaf Blades:

The - N treatm en t produced the s m a lle s t amount of

le a f grow th (1 3 .2 g), and - F e, 5 Mo, and the check, in d escen d in g ord er, had
the la r g e s t amount of le a f grow th (37. 65, 37. 50 and 37. 15 g r e sp e c tiv e ly ).
The sam e gen era l trend o b serv ed for the stem and p e tio le dry w eight
accu m ulation w as found for le a f growth.

Nine of 11 e lem en ts produced le s s

le a f grow th at both the low and high le v e ls than found for the ch eck treatm en t.
In crea sin g le v e ls of iron again d e c re a sed le a f dry w eight and in c r e a sin g le v e ls
of m olybdenum in c r e a se d it.
No s ta tis tic a l d iffe re n c e s could be found betw een any m inor elem en t
treatm en t and the check.

The - N, - K, 5 K (Cl), 5 Ca (SO^), and 5 Mg

treatm en ts r e su lte d in sig n ifica n tly l e s s le a f growth than the check treatm ent.
The 5 K (SO .) trea tm en t did not r e su lt in a sign ifican t reduction of le a f growth.
The 5 M g treatm en t produced sig n ifica n tly le s s le a f growth than the - Mg
treatm en t.

ROOTS
1

I 1 I I— I i— I— r

i— i— i— i

LEAF BLADES

PETIOLES

STEMS
t—

T

r

J

I

1

I

1

I

|

I

|

Check

m
■

i*
5K(S04)
5K (Cl)
-C a
5Ca(S04) P
5Ca(CI)

I
i*

■

I
i
i
-M o

j

10
LSD

I I

I i

L

10 20 3 0
0 5
10 2 0 3 0
20 30 40
DRY WEIGHT ACCUMULATION - (percent o f to ta l dry w eig h t)

5%
I%
Slg. different _ * a t 5 %
than check
**a t I%

40

50

A ccu m ulation of Dry W eight E x p r e sse d in Per Cent of T otal D ry Weight by
R oots, Stem s, P e tio le s, and L ea f Blades (s e e figu re opp osite)
A co m p a riso n of dry w eight accu m ulation o f the individual plant p arts
m ade it p o s s ib le to evalu ate the e ffe ct of the trea tm en ts upon ro o ts, stem s,
p e tio le s and le a f b la d es.

By con v ertin g th ese v a lu es into p er cent of the total

dry w eight accu m u lated by a plant, the influence of the trea tm en ts on sp e c ific
plant p arts in rela tio n to the en tire plant w as dem onstrated.
Roots:

The ro o ts w ere 26. 29% of the total dry w eight produced with

" 1’ ’

1

the ch eck treatm en t.

No treatm en t had a sig n ifica n tly low er p ercen ta g e of

roots than the check.

The - N, - P, - Cu, 5 Mg, - Mo, and 5 Mo treatm en ts

had a sig n ifica n tly la r g e r p ercen ta g e of roots than the check.
By com paring high and low le v e ls of the individual e lem en ts, only
nitrogen, phosphorus, and m agnesium treatm en ts w ere different sta tis tic a lly .
The - N, - P, and 5 M g trea tm en ts w ere la rg er at the 1% le v e l than 5 N, 5 P
and - M g r e s p e c tiv e ly .

With in c r e a sin g nitrogen and phosphorus, or d e c r e a s ­

ing m agnesium in the solution, the p ercen ta g e of roots d e c re a sed .

A g rea ter

\

p ercen ta g e of r o o ts w as produced in both low and high le v e ls of a ll other e le
m en ts than found for the ch eck treatm ent.
Stem s:

No treatm en t w as sig n ifica n tly la rg er in p ercen ta g e of stem

grow th than the check.

The - N, - P, - K, - Ca, 5 Ca(Cl), 5 Mg, and - Mo

w ere s m a lle r in p ercen ta g e of stem growth than the check.

By com paring

52.

high and low le v e ls o f s p e c ific e le m e n ts, the - P treatm en t had sig n ifica n tly
l e s s stem grow th than the 5 P treatm ent, w hile the - M g treatm en t had s ig ­
n ifica n tly l e s s than the 5 M g treatm en t.
P e tio le s:

T h ree trea tm en ts, - K, 5 Ca (Cl), and 5 M g w ere s ig n i­

fican tly s m a lle r in p e r ce n ta g e of p e tio le s than the check, but no treatm ent
w as la r g e r than the check .

The - K treatm ent had sig n ifica n tly le s s p e r ­

cen tage of p e tio le grow th than the 5 K (SO J and the 5 K (Cl) treatm ents*
4
A lso 5 M g had sig n ifica n tly le s s p ercen ta g e of p etio le growth than the - Mg
treatm en t.
L eaf Blades:

The m o st pronounced e ffe c t of any treatm ent was

e x p r e s s e d by 5 N, w hich produced 30% m ore le a f b lad es than - N.

While

5 N w as the only treatm en t having sig n ifica n tly m ore le a f grow th on p e r ­
cen tage b a s is than the check , - Mn, - Cu, and 5 Mo had sig n ifica n tly
le s s than the check .

By com paring high and low le v e ls of sp e c ific e l e ­

m en ts, - P and 5 M g had l e s s le a f grow th on a p ercen ta g e b a s is than 5 P
and - M g r e s p e c tiv e ly .
Shoot/R oot R atio:

By adding the dry w eight of the above-ground

p a rts, and dividing them by the root dry w eight, the follow ing sh o o t/r o o t
ra tio s w e r e esta b lish ed :

T reatm en ts
5 P, - Mg
5 N
5 K (Cl)
Check
5 K (SO .)
- Fe
- K, 5 C a(S04), 5 F e
5 B
- B, - Ca
5 Ca (Cl), - Mn, 5 Mn, 5 Cu, - Zn
5 Zn
- Mo
- N, - P, - Cu, 5 Mo
5 Mg

Ratio
3. 9
3. 7
3. 1
2. 8
2. 6
2. 2
2. 1
1. 9
1. 8
1 .7
1.6
1 .4
1. 3
1. 2

In creasin g le v e ls of nitrogen, phosphorus and p otassiu m and d e c re a sin g
le v e ls of m agn esium in the solution a lso in c r ea se d the sh o o t/r o o t ratio.
O therw ise, the high and low le v e ls of calcium and a ll m inor elem en ts p r o ­
duced low er s h o o t/r o o t r a tio s than the check.

W E IG H T

NITROGEN LEVEL IN SOLUTION G 2E3 4 2 PPM
2tO PPM (CHECK)
1050 PPM

K
S2 200

DRY

£

PERCENT

o
J

150

CONTENT

-

0.

S 100

o

NUTRIENT

o

!

S

1

Co

m
Mg
PETIOLE

m
Cu

Fe

Zn

COMPOSITION

5.2
5.0,
2.4
WEIGHT

—
—
•w
*

NITROGEN

CONTENT

- PERCENT

DRY

20

PETIOLES
STEMS
CHECK PETIOLES 8 STEMS
NOT ANALYSED

K K
(S04 )(C I)

Mg
Co Ca
(S04)(CI)
SOLUTION CONTENT

Mn

Cu

Zn

Mo

54.

NUTRIENT ELEM ENT COMPOSITION OF PETIOLES AND STEMS

Data p erta in in g to the p e tio le and stem co m p o sitio n for plants trea ted
w ith the v a r io u s nutrient so lu tion s are given in Appendix T a b les 7 and 8.

T h ese

ta b les and the follow in g graphic illu str a tio n s and d isc u ssio n show the effect on
p e tio le co m p o sitio n when one s p e c ific elem en t w as v a ried and a il other elem en ts
kept at a constant le v e l, a s r e p r e se n te d in the ch eck treatm ent and the influence
of v a ry in g the supply of d ifferen t e lem en ts upon p e tio le com p osition of a s p e c i­
fic elem en t.

N itrogen (s e e figu re opp osite)
With in c r e a sin g le v e ls of nitrogen in the solution (42, 210 - check, and
1050 ppm) nitrogen, phosphorus, boron and copper concen tration of the p e tio le s
in c r e a se d and calciu m content of the p e tio le s d ecrea sed .

P otassium , m a g n es­

ium and zin c w e r e found to be h igh est w ith 210 ppm nitrogen, but d ecrea sed
w ith e ith e r in c r e a se d or d e c r e a se d le v e ls of nitrogen in the solution.

Iron

and m an gan ese, on the oth er hand, show ed an in v e r se relation sh ip and in ­
c r e a s e d with e ith er in c r e a s e d or d e c re a sed nitrogen le v e ls , and w ere low est
w ith 210 ppm n itrogen in the solution .

A five fold in c r e a se of nitrogen in

the solu tion in c r e a se d the p e tio le n itrogen fiv e tim e s that of the check tr e a t­
m ent, doubled phosphorus, trip led boron and r a ise d m an ganese from 63 ppm
for the ch eck treatm en t to 159 ppm .

At the sam e tim e, p otassiu m w as

55.

d e c r e a se d 60% and ca lciu m 50% by in c r e a sin g nitrogen to 5 N.
The low er graph show s that nitrogen in the solution had the la r g e s t
e ffe ct on n itrogen in stem and p e tio le s .

Although the nitrogen content w as

low er in the p e tio le s than in the s te m s of the - N treatm ent, both stem
and p e tio le n itrogen content w ere n early alik e w ith 210 ppm nitrogen in
the solu tion .

The 5 N treatm en t show ed 3% m o re actual nitrogen in the

p e tio le s than in the s te m s.

In the - P treatm ent, nitrogen in p e tio le s and

s te m s w as a ls o v e r y low and in c r e a se d with in c r ea sin g phosphorus in the
solution .

T he only oth er treatm en t w hich see m e d to sig n ifica n tly a ffect

n itrogen w as - Ca, w h ere both stem and p e tio le nitrogen w ere v e ry high.
An in c r e a s e in the supply of p otassiu m and boron see m e d to reduce the
amount of n itrogen in both stem and p e tio le s .

An in c r e a se in the m agnesium

and iro n supply appeared to in c r e a s e nitrogen in both stem and p e tio le s.

W E IG H T

PHOSPHORUS LEVEL IN SOLUTION E iiS 3 O PPM
31 PPM (CHECK)
155 PPM

PERCENT

DRY

(5 iuu
£
>tk
ca
5

0.

150

C O N TE N T-

§
oo

100

I

NUTRIENT

ft
50

in

2

v M

Co

PETIOLE

Mn

Cu

Zn

COMPOSITION

1.0

PETIOLES
STEMS
CHECK PETIOLES
CHECK STEMS

PHOSPHORUS

CONTENT-

PERCENT

DRY

WEIGHT

Fe

Mg

N

5 -P
N

5

-K
P

5
5 -C o 5
5
K
K
Ca Cq
<S04 XCI)
{SO4 MCI)

5 -F e
Mg

5
Fe

Mn 5
Mn

SOLUTION CONTENT

5
Cu

Zn

Mo

56 .

Phosphorus (s e e figu re opp osite)
An in c r e a s e in the supply of phosphorus in the solution (0, 31 - check,
and 155 ppm ), in c r e a se d the am ount of ph osphorus, nitrogen, boron, and
copper in the p e tio le s and d e c r e a se d the calcium content.

Both potassium

and m agn esium w ere h igh est w ith 31 ppm phosphorus in the solution, but
both d e c r e a se d w ith e ith e r in c r e a se d or d e c re a sed le v e ls .

M anganese

show ed a r e v e r s e relation sh ip with 31 ppm of phosphorus in the solution
r esu ltin g in the lo w est m an ganese content.
When phosphorus in the solution w as in c r ea se d to five tim es the
control, the phosphorus content of the p e tio le s w as r a ise d th ree fold and
boron four fold that found in the con trol treatm ent, but calcium w as d e ­
c r e a se d to 50% of the check.

M anganese was m ost s e v e r e ly affected when

the solu tion phosphorus w as in c r e a se d from 0 to 31 ppm .

In th is ca se,

m an ganese in the p e tio le s w as low ered from 247 to 63 ppm.
A com p arison of stem and p etio le com p osition is shown in the low er
graph.

The stem phosphorus content w as a little higher than the p etio le

content w ith 0 phosphorus in the solution.
sym ptom s w ere found.

H ere again, v isu a l d eficien cy

T he d ifferen ce betw een stem and p etio le content

b ecam e la r g e r with in c r e a sin g phosphorus in the solution.

At the 31 ppm

ph osphorus le v el, it com pared . 26 and . 37%, and at the 5 P le v el . 40 and
1. 02% w ith the p e tio le s being higher in both c a s e s .

57.

C o n sid erab le in flu en ce on phosphorus content in the p e tio le s w as
ex erted by the follow in g treatm en ts:
5 Mg.

5 N, the th ree K trea tm en ts, - Ca and

In a ll c a s e s , the p e tio le phosphorus content w as sig n ifica n tly higher

than that found for the ch eck treatm en t.

The - N treatm en t had a phosphorus

p e tio le content that w as sig n ifica n tly low er than the one found in the check
treatm ent.

T he stem phosphorus content w as not sig n ifica n tly different

from the ch eck for any of the trea tm en ts.

POTASSIUM LEVEL IN SOLUTION
t-m sa o PPM
IT O 1 235PPM (CHECK)
■ S B 1171 PPM K(S04 >
1171 PPM K(CI)
NOT ANALYZED

10

WEIGHT

250

PERCENT

DRY

200

CONTENT

-

150

NUTRIENT

8

100

£
£
50

m
Co

Mg

Fe

Mn

Cu

PETIOLE COMPOSITION

POTASSIUM

CONTENT

PERCENT

DRY

WEIGHT

PETIOLES
STEMS
CHECK PETIOLES
CHECK STEMS

SOLUTION

CONTENT

Zn

58.

P otassiu m (s e e figu re opp osite)
In c re a sin g the le v e l of p o ta ssiu m in the nutrient solution (0, 234 - check,
1171 ppm) in c r e a s e d the p o tassiu m , boron, and zin c content of the p e tio le s, but
d e c r e a se d m agn esium and m an gan ese.

The high le v e l of p otassiu m w as tested

w ith both su lfate and ch lorid e a s anions.

With p otassiu m sulfate, 1% m ore

actual p o ta ssiu m w as found in the p e tio le s than with potassium ch lorid e.
Boron d e c r e a se d from 229 to 198 ppm and phosphorus w as a lso sig n ifica n tly
low er when p o ta ssiu m ch lorid e in stead of potassiu m sulfate w as used.

With

0 le v e l of p o ta ssiu m in the solution, the lo w est p etio le copper and zin c co n ­
tent of any treatm en t a s w ell a s the h igh est m agnesium content w ere p r o ­
duced.

The iron content of p e tio le s w as a lso com p aratively high.

When

p otassiu m w as in c r e a se d from z e r o to 234 ppm in the solution, the m anganese
content dropped from 268 to 63 ppm.

The p otassiu m p etio le content of 5. 72%

w as found when 234 ppm p otassiu m w ere used in the check solution.

T his

w as n ea rly th ree fold the amount found in a norm al field sam ple.
In the low er graph, a com p arison of stem w ith p etio le potassium
content for the - K treatm en t showed that the ste m s had a higher potassium
content than the p e tio le s .

When the p otassiu m content of the solution w as

in c r ea se d , the p otassiu m content of the p e tio le s w as g rea ter than the stem
content.

With 234 ppm p otassiu m in the solution, the ste m s showed 2. 37%

and the p e tio le s 5. 72% p o ta ssiu m .

T his d ifferen ce becam e even la rg er

with 1171 ppm in the solution w here with the u se of potassium sulfate the
stem s accum ulated 2. 61% and the p etioles 9. 86% potassium , as compared
to 2. 68 and 8. 88% potassium resp ectively, when potassium chloride was
used.
Furtherm ore, a s shown in the lower graph, the potassium content
of the p etio les for the - N, 5 N, - P, 5 P, - K, 5 Ca, - Mg, 5 Mg, - Fe, 5 Fe,
- B, - Cu, 5 Cu, - Mo and 5 Mo treatm ents was below that found for the check
treatm ent.

A potassium content higher than that found in the check trea t­

m ent was in p etio les from 5 K, - Ca, and 5 Zn treatm ents.
*
In a ll treatm ents, stem an alysis for potassium was below that of the
check treatm ent, except in the 5 P and 5 K treatm ents.

The difference b e­

tween p etio le and stem a n alysis for potassium was le s s for the 5 N than - N,
5 P than - P, 5 Ca than - Ca, 5 Mg than - Mg, and 5 B than - B treatm ents.
Petiole a n alysis showed that the potassium petiole content in th ese 5x treat­
m ents w as low er than in their resp ectiv e minus treatm ents, but a rev erse
relationship of potassium in the stem s, as indicated by stem analysis, was
found.

A lso the 5 Ca (SO.) treatm ent resulted in le s s reduction of potas-

sium content in both p etio le and stem tissu e than the 5 Ca (Cl) treatm ent.

WEIGHT

CALCIUM LEVEL IN SOLUTION
0
PPM
200 PPM (CHECK)
1002 PPM Co(S04)
1002 PPM Co(CI)

150

CONTENT -

PERCENT

DRY

u

NUTRIENT

100-

Mg

Fe

PETIOLE COMPOSITION
260
PETIOLES
STEMS
CHECK PETIOLES
CHECK STEMS

L80

1.40

CONTENT

-

PERCENT

DRY

WEIGHT

220

LOO

.6 0

.2 0

(S04) (Cl)

SOLUTION CONTENT

Calcium (see figure opposite)
With in crea sin g le v e ls of calcium in the solution (0, 200 - check, and
1002 ppm), calcium in crea sed in the p etioles, while potassium , m agnesium,
and boron d ecreased .

Nitrogen and phosphorus content of the p etioles w ere

higher with - Ca than 5 Ca.

Iron and m anganese w ere low est in the check

treatm ent, and in crea sed with either in creased or d ecreased lev els.

Minor

variations in the copper and zinc content of p etioles w ere associated with a
variable calcium supply.

The high lev el of calcium was tested with both

sulfate and chloride as anions.

A highly significant in crease of calcium in

the p etio les w as observed when calcium chloride instead of calcium su l­
fate w as used (2. 55 and . 96%).

At the sam e tim e, there was a decrease

in potassium (from 4. 51 to 2.82%), m anganese (from 131 to 76 ppm), and
boron (from 67 to 37 ppm) when the calcium chloride was used instead of
calcium sulfate.
Comparing stem and p etiole an alysis for calcium , as shown in the
lower graph, the calcium content for both plant parts was nearly the same
at the 0 lev el.

This w as found also w here calcium sulfate was used, but

with calcium chloride, the petiole content was 1% higher than the stem
content.

Although there w as a m arked in crease in both stem s and p etioles,

none of th ese treatm ents produced a higher calcium content of stem s than
in p etio le s.

However, in the 5 N, 5 P and 5 Mg treatm ents, the calcium

61.

in ste m s and p e tio le s w as v e r y c lo s e .

Sign ifican tly low er calcium p etio le

content than in the ch e ck w as found in the follow ing trea tm en ts: 5 N, 5 P
and 5 Mg, w h ile - N, - P, - Mg, - Fe, 5 Fe, 5 B , - Cu, - Mo and 5 Mo
w ere sig n ifica n tly high er than the check.

In the ste m s, only the - Mg

treatm ent had a sig n ifica n tly higher calciu m content than the check.

l e v e l IN s o l u t o n

W E IG H T

m a g n e siu m

^

^

PERCENT

CONTENT
NUTRIENT

2b

L
i
j

i

B
Ca
PETIOLE

COMPOSITION

IOOi

MAGNESIUM

CONTENT

- PERCENT

DRY

WEIGHT

PETIOLES
STEMS
fr™- CHECK PETIOLES
CHECK STEMS

K K
(S04) (Cl)

Fe
Ca Co
(S04)(CI)
SOLUTION CONTENT

Mn

Cu

Zn

CHECK)

2 4 3 PPM
NOT ANALYZED

DRY

*

™

Mo

62.

M agnesium (se e figu re opp osite)
When the le v e l of m agn esium in the solution w as in c r ea se d (0, 49 check, and 243 ppm), m agn esium and boron in c r ea se d in the p e tio le s, w h e r e ­
as calciu m , copper and z in c d e c r e a se d .

Phosphorus, iron and m anganese

w ere lo w est w ith 49 ppm m agn esium in the solution, and w ere in crea sed
when the le v e l of m agn esium in the solution w as e ith er in c r ea se d or d e­
c r e a se d .

P otassium on the other hand, w as high est with 49 ppm m agnesium

in the solution, and d e c r e a se d w ith e ith er an in c r e a se or d e c r e a se in the
m agnesium supply.

N itrogen showed a d e c r e a se in the p etio le when m a g ­

nesium w as in c r e a se d from 0 to 49 ppm.

(P etio les of the 5 M g treatm ent

w ere not an alyzed for nitrogen b ecau se of sh ortage of p e tio le s).
A fiv e fold in c r e a s e of m agnesium in the solution n ea rly doubled the
p etio le m agn esium and phosphorus concentration.

F u rtherm ore, m anganese

w as in c r ea se d sig n ifica n tly from 63 to 174 ppm and boron from 56 to 152
ppm.
In the co m p a riso n of stem w ith p e tio le com position , a s shown in
the low er graph, th ere w as a gen era l tendency for the stem m agnesium to
be low er than the p e tio le m agn esiu m .

T his com p arison w as true for all

treatm en ts excep t for the 5 K (SO4), 5 K (Cl) and - Mg trea tm en ts.

With

th ese trea tm en ts, the stem m agnesium w as higher than the p etio le m a g ­
n esiu m .

In a ll th ree c a s e s , v isu a l d eficien cy sym ptom s on the le a v es w ere

63.

o b serv ed .

The m o st pronounced e ffe ct on m agn esium uptake w as not ex erted

by the m agn esium w ithin the solution, but by the p o ta ssiu m .

H ere, the - K

treatm en t produced a m agn esiu m content in the p e tio le s of . 99%, which
com pared to the 74% M g of the 5 Mg treatm ent.
Significant in c r e a s e s o f p e tio le m agnesium w ere found in the - K and
- Fe trea tm en ts, w hile - N, - P, 5 K (SO^), and 5 K (Cl) w ere sign ifican tly
low er in p e tio le m agn esium than the check.

NUTRIENT

CONTENT

- PERCENT

DRY

W EIG H T

IRON LEVEL IN SOLUTION ESS23 O PPM
IT T H 5 PPM (CHECK)
2 5 PPM

m
Cu
PETIOLE

Zn

COMPOSITION

00
PETIOLES
STEMS
? CHECK PETIOLES
CHECK STEMS

60

IRON

CONTENT

-

PPM

DRY

WEIGHT

80

40

"7

20

K K
(S04 )(CI)

Mn

Ca Ca
(S04) (Cl)
SO L U T IO N

CONTENT

Cu

Zn

Mo

64.

Iron (s e e fig u re opp osite)
When the le v e l of iron w as in c r e a se d in the solution (0, 5 - check, and
25 ppm), v e ry little e ffe ct upon iron in the p e tio le s could be o b served .

M ost

of the e le m e n ts show ed no sign ifica n t in c r e a s e or d e c r e a se in the p e tio le s .
When iron in the solu tion w as in c r e a se d from 5 to 25 ppm, calcium in crea sed
sign ifican tly, w h erea s p o ta ssiu m d e c re a sed in the p e tio le s.

An in c r e a se of

iron from the 0 le v e l to 5 ppm in the solution brought about a sign ifican t d e­
c r e a s e of m agn esium in the p e tio le s from 65 ppm to 45 ppm.

A general te n ­

dency p r e v a ile d for 5 ppm iron to r esu lt in e ith er the h igh est or the low est
value in the p e tio le for other e lem en ts w hich in c r ea se d or d ecrea sed when
iron w as e ith e r in c r e a se d or d e c re a sed in the solution.
From the com p arison of the stem and p e tio le iron content, a s shown in
the low er graph, it can be o b serv ed that the iron in the solution has v ery little
effect upon iron in both ste m s and p e tio le s .

In a ll th ree iron treatm en ts, the

stem iron w as higher than the p e tio le iron, which actually w as the c a se in m ost
other trea tm en ts.

H ow ever, an in te r e stin g exception to this w as the 5 Mn

treatm ent.

H ere, the p e tio le iron content w as 90 ppm com pared to 50 ppm in

the s te m s .

A lso th ere w as a g r ea ter amount of iron in the p e tio le s than in

the stem for the - M g and - Zn trea tm en ts.

On the other hand, the - K tr e a t­

m ent had 100 ppm iron in the stem and 70 ppm in the p e tio le s .

The - Ca

treatm en t had a s im ila r relation sh ip with 80 ppm iron in the stem and 50
ppm in the p e tio le s .

O th erw ise, the iron v a lu e s for both p e tio le s and

ste m s w ere c lo s e ly grouped w ith the excep tion of the above m entioned
treatm en ts, and no sig n ific a n c e w as found in the a n a ly sis of variance for
iron in e ith er of the two plant p a rts.

MANGANESE LEVEL IN SOLUTION E 2 S 3 0 PPM
17773 .5 PPM (CHECK)
2 5 PPM
300

6r

oX

G 5

£

v> 2 5 0
£

>.
<
fc
Q

>-

§

sU
oCtj 4
fe!

200

3o.
0.
150

fc!

I
§ 2
%
UJ
tfc

oo

100

s
£
vC-

i

Co

Mg
PETIOLE

I J|»

50

Fe

M

Mn

W EIGHT

I

Cu

PETIOLES
STEMS
flaw? CHECK PETIOLES
CHECK STEMS

250

CONTENT

— PPM

200

150

100
MANGANESE

Zn

COMPOSITION

300

DRY

12

50

N

5 -P
N

5

P

-K

5
5 -C a 5 5
K
K
Co Ca
<S04)(CI)
(S04)(CI)

5 -F e 5
Mg
Fe

Mn 5 -B
Mn

SOLUTION CONTENT

5
B

5
Cu

Zn

66.

M anganese (s e e figu re opp osite)
In crea sin g le v e ls of m an gan ese in the solution (0, . 5 - check, and
2. 5 ppm) brought a sig n ifica n t change of m an ganese in the p e tio le s.
oth er e le m e n ts excep t iron w ere a ffected , but v e ry little .

The

Iron in the

p e tio le s in c r e a se d from 35 ppm in the - Mn treatm ent to 90 ppm when
high le v e ls of m an gan ese w ere used in the solution.
The low er graph show s a com p arison betw een stem and p etiole
content of m an gan ese.

Although the d ifferen ce betw een stem and p etiole

m an ganese content in c r e a se d with in c r ea sin g le v e ls of m anganese in the
solution, the - Mn treatm en t show ed a higher m anganese concentration
in the ste m s than in the p e tio le s .

H ere, too, v isu a l d eficien cy sym ptom s

on le a v e s w ere o b serv ed .
Other e lem en ts e x er te d a g r e a te r influence upon m anganese up­
take than m an gan ese upon oth er e lem en ts.

If the check is taken as a

b a sis of com p arison , the follow in g trea tm en ts sig n ifica n tly in crea sed
m anganese in the p e tio le s:

5 N, - P, - K, - Ca and 5 Mg.

In gen eral, a ll m an gan ese p etio le v a lu es w ere rather low in
com p arison with a v era g e fie ld sa m p le s.

T his w as a lso ob serv ed for the

high le v e l of m an gan ese with 2. 5 ppm of m an ganese in the solution.

BORON l e v e l in

PETIOLE

SOLUTION ESSSSl o PPM
■m
E Z 2 3 5 PPM (CHECK)
11
Bffiffl 2 5 PPM

COMPOSITION
PETIOLES
STEMS
CHECK PETIOLES
CHECK STEMS

240

k. 200

£
£

160-

^

120

Cl

s
l

k.

C> 8 0

**

40

5 -Cu
K K
(S04)(C I)

Co Ca
(S04 )(CI)
SOLUTION CONTENT

Mn

■Zn
Cu

-M o
Zn

Mo

67.

Boron (s e e fig u re op p osite)
In crea sin g the le v e l of boron in the solution (0, . 5 - check, and 2. 5
ppm) brought about a sig n ifica n t in c r e a se in the boron and calcium content
of the p e tio le s and a sig n ifica n t d e c r e a se in phosphorus and copper co n ­
c en tra tio n s.

A fiv e fold in c r e a s e o f boron in the solution resu lted in a

three fold in c r e a s e in the p e tio le boron.

P otassium w as high est with

. 5 ppm of boron in the solu tion and d e c re a sed with e ith e r low er or higher
le v e ls of boron.

N itrogen, m agnesium , iron and m an ganese had a r e ­

v e r s e rela tio n sh ip and in c r e a se d with eith er low or high le v e ls of boron.
By com paring the p e tio le and stem a n a ly sis for boron, as shown
in the low er graph, the d ifferen ce betw een stem and p etio le, boron co n ­
cen tration in c r e a se d w ith in c r e a sin g boron in the solution.

With high

le v e ls of a ll n u trien ts the p e tio le content for boron w as alw ays higher
than the stem content.

In th ree treatm en ts (-M n , - Cu, -M o), the stem

boron content w as h igh er than the p etio le content.

The graph a lso show s

that the boron content w as influenced m o re by the variation of other e l e ­
m ents in the solution than by boron its e lf.

The -P treatm ent produced

nearly the sam e low v a lu e s for boron in p e tio le s and ste m s as the - B
treatm en t.

The 5 P treatm en t produced the h igh est boron p etio le content

for a ll trea tm en ts.

T rea tm en ts other than 5 B that had a sig n ifica n tly

higher p e tio le boron content than the check treatm ent w ere 5 N, 5 P,
5 K(S04), 5 K(C1), - Ca, 5 M g and 5 Zn.

COPPER LEVEL IN SOLUTION

0 PPM
0 2 PPM (CHECK)
10 PPM

60

P50

K
§
^ 5
§

I

LI

g 3
i

o
£2
Uj

E

£
>§
S 40
0.

i

1
IIp
i
IH

g

^

10

i |il
ig *1 a
Fe

Mg

PETIOLE

„

!|i
li 1
1
l |
!g
i

■

Co

!

20

i

iis s

&

I

30

K
oS
o
!z
UJ
£
h-

i# ^!S
Mn

Cu

Zn

COMPOSITION

40

30

I
>-

0:
Q

(—
■

20

§
PETIOLES
STEMS
CHECK PETIOLES
CHECK STEMS

P=
£

8

K K
(S04)(CI)

Co Ca
(S04)(CI)

Mg

SOLUTION CONTENT

Mn

Cu

Zn

Mo

68.

Copper (s e e fig u re op p osite)
In crea sin g le v e ls of copper in the solution (0, . 02 - check, and
, 1 ppm) had no sig n ifica n t e ffe ct on any elem en t excep t calciu m .

In both

0 and . 1 ppm copper le v e ls , calcium w as sig n ifica n tly higher than in the
check treatm en t.

The other m ajor elem en ts w ere affected v ery little .

While iron and m an gan ese w ere h igh est with 0 copper in the solution,
boron w as h ig h est with the . 02 ppm le v e l of copper w h ereas copper
and zin c w ere h igh est with . 1 ppm copper in the solution.
The low er graph show s a com p arison betw een stem and
p etio le copper concen tration.

It w as obvious that a large number of

treatm en ts resu lted in higher copper contents in the ste m s than in the
p e tio les; how ever, the th ree trea tm en ts varyin g in copper content of
the nutrient solu tion w ere higher in p etio le copper than in stem copper.
Other trea tm en ts w hich follow ed th is m ore norm al trend w ere 5 K(SO^),
5 Ca(S04>, - Mg, - F e, - Mn, - B and 5 Zn.

The lo w est copper content

of p e tio le s w as found in the p e tio le s of the - K treatm ent, w hile the - Cu
treatm en t produced a high copper content.

Both stem and p etio le a n a ly sis

for copper show ed no sig n ifica n ce betw een treatm en ts, w hich w as a lso o b ­
ser v e d in the a n a ly s is of varian ce for iron.

The low p etio le copper values,

how ever, in the - K and 5 Ca(Cl) trea tm en ts a s w ell a s the high stem copper
content of the treatm en t 5 Mo m erit attention.

ZtNC LEVEL IN SOLUTION E S 3

0 PPM
.05 PPM (CHECK)
.25 PPM

6W EIG H T

yA

DRY

a: 100

-

PERCENT

a

80 -

NUTRIENT

CONTENT

m

m.

11

Hi

m .

Co

Mg
PETIOLE

Fe
COMPOSITION

60

50

>.

40

■WH- /
30

20

=> ------ PETIOLES
$
STEMS
# — - CHECK PETIOLES
| •*— CHECK STEMS

&

W
N

5 -P
N

5 -K
P

5
5 -C o 5
5
K K
Co Co
(S 04)(CI)
(S04)(CI)

5 -F e 5 -Mn 5 - 8
Wig
Fe
Mn

SOLUTION CONTENT

5
B

m

-m

■

if i

Cu 5 ■Zn 5 -M
Cu
Zn

Mo

. Zinc (see figure opposite)
An in c r e a s e in the le v e ls of zin c in the nutrient solu tion (0, . 05 check, and . 25 ppm) r e s u lte d in an in c r e a s e in p o ta ssiu m and boron in
the p e tio le s , but d e c r e a se d m an gan ese and z in c.

The only elem en t w hich

w as sig n ific a n tly a ffected w as boron.
A s shown in the low er graph, 23 of the 25 treatm en ts had a higher
zin c content in the p e tio le s than in the ste m s.

Only - K and 5 Mo had

low er zin c con cen tration s in the p e tio le s than in the ste m s.

The highest

z in c p e tio le content w as found in the - Mo and 5 Ca(Cl) trea tm en ts, w hile
the lo w e st content w a s o b se rv e d in the - K treatm en t.

The lo w est zin c

stem content w as found in the - N treatm en t and the h igh est in the - Mg
treatm en t.
Z in c stem v a lu e s show ed sig n ifica n t d ifferen ces but not the
p e tio le v a lu e s.

When m agn esium in the solution w as in c r ea se d from 0 to
('

243 ppm, the z in c content o f ste m s d e c r e a se d sig n ifica n tly .

On the other

hand, in c r e a s e d le v e ls of boron in the solu tion (from O to 2. 5 ppm) in ­
c r e a s e d sig n ifica n tly the z in c content of stem s.

k-

2 .5

£

2.0

K-

PETIOLES

1.0

STEMS

u

.49 .55

4.42

7.37

O

2 8 4 532

849
192

1418
192

5 K (Cl)

5Ca(CI)

Treatment Ck 5Mg 5Ca(S04 )

C l/S 0 4 RATIO IN SOLUTION

Chlorine (see figure opposite)
A s m en tion ed e a r lie r , high le v e ls of p o ta ssiu m and calcium w ere
te ste d w ith s u lf a t e - -5 K (SC>4) and 5 Ca(SC>4) tr e a tm e n ts--a n d w ith c h lo r ­
i d e - -5 K(C1) and 5 C a(C l) tr e a tm e n ts --a s anion s.
w as added to the high m agn esiu m (5 Mg) treatm en t.

Som e ch lorid e a lso
The graph opposite

show s the c h lo rid e con cen tration of ste m s and p e tio le s in com p arison
w ith the Cl_ ra tio in the so lu tio n s of the check, 5 Mg, 5 C a(S04), 5 K (Cl)

so4

and 5 Ca(Cl) tr e a tm e n ts.
p etio le ch lorid e con ten ts.
%

In crea sin g r a tio s brought in c r e a sin g stem and
The ste m s, how ever, did not in c r e a se in

chloride a s m uch a s the p e tio le s and the d ifferen ce betw een th ese two
plant p a rts b ecam e la r g e r with in c r e a sin g c h lo r id e /su lfa te r a tio s in the
solution.
The ch lorid e content o f p e tio le s from the ch eck treatm en t w as
. 07%.

C hloride v a lu e s of 1. 70% for the 5 K(C1) treatm en t and 1. 89%

for the 5 Ca(Cl) treatm en t w e re in the approxim ate range o f to x icity .
H ow ever, no ch lorid e to x ic ity sum ptom s on le a v e s w ere o b serv ed .

72.

DISCUSSION

T he c o m p o sitio n of the nutrient so lu tio n s u sed in th is exp erim en t
seem ed to be the m ain fa cto r in flu en cin g grow th and developm oit of the
plant.

Bonner and G a lsto n e (1952) d e scr ib e d grow th a s quantitative m atter

w hich can be m e a su r e d (length and w eight) w hile developm ent is a qualita­
tive e x p r e s s io n and can only be o b se rv e d .

The r e s u lts , th erefo re, should

be broken down into dry w eigh t accu m ulation and length of shoots a s growth,
w hile v isu a l d e fic ie n c y or to x ic ity sym p tom s a s w e ll a s va ria tio n s of lea f
c h a r a c te r is tic s should be tr e a ted a s developm ent.

H ow ever, both grow th

and developm ent e x p r e s s th e m se lv e s a s an entity and w ill be trea ted a s
such.

Shear, Crane and M e y e r s 1 (1946) concep t of nutrient balance and

in ten sity can be taken a s a b a sis fo r evaluation of the so lu tio n s in r e sp e c t
to dry w eight accu m u lation and lin ea r grow th.

T h ese authors fe lt that a

m ore id eal balance b etw een the e s s e n tia l nutrient e lem en ts has to e x is t
before h igh er y ie ld s can be obtained by in ten sify in g nutrition.
P r e lim in a ry ex p e rim en ts show ed that the lx le v e l of the No. 1
solution p ro p o sed by Hoagland and Arnon (1950) m ight be suitab le a s a
standard solu tion and a ll other trea tm en ts w e re adjusted accordin gly.

A

com p arison of the p e tio le co m p ositio n of th is ch eck treatm en t w ith the
standard p e tio le co m p o sitio n for M ichigan Concord grap es, a s esta b lish e d

by L a r se n (1957) o v e r a num ber o f y e a r s , r ev e a le d som e m arked d iffe r ­
en ces.

T ab le 2 show s th is co m p a riso n .

TABLE 2
COMPARISON BETWEEN STANDARD PETIOLE VALUES OF FIELD SAMPLES
(LARSEN, 1957) AND VALUES OF THE CHECK TREATM ENT

E lem en t

N itrogen
Phosphorus
P otassium
Calcium
M agnesium
Iron
M anganese
Boron
Copper
Z inc

%
%
%

7o
7c
ppm
ppm
ppm
ppm
ppm

F ie ld Standard
V alu es

N utrition Solution
Check T reatm ent V alues

.8 0
.2 4
1. 91
1.60
. 46
44
434
25
35
23

1 .0 6
. 37
5. 72
.99
.4 5
35
63
56
24
46

'

Inspection of th e se v a lu e s show s that p otassiu m in p e tio le s o f the
check treatm en t w as th ree tim e s high er than the standard fie ld a v era g e.
C alcium , on the o th er hand, w a s 50% low er, w hile boron and z in c showed
double the amount found under fie ld con d ition s.

The la r g e st variation,

how ever, w as for m an gan ese w here the fie ld sa m p le w as sev en tim e s
higher than that for the nutrient solution sa m p le.

The resp o n sib ility for

such a m arked v a ria tio n could lie in the follow in g p o s s ib ilitie s :

1. T he solu tion is e ith e r too high or too low in the sp e c ific
elem en t.
2. In teraction o f oth er e lem en ts m ight p rev en t o r enhance
uptake of the s p e c ific elem en t.
3. L uxury consum p tion m ight o ccu r w here ex tr em e high le v e ls
a r e p r e se n t.
4. The e lem en t m ight be tied up in the solution and not a v a il­
ab le to the plant.
Before th e s e p o s s ib ilit ie s can be d efin itely identified, oth er fa c to r s,
such a s p e tio le co m p o sitio n of oth er trea tm en ts, a s w e ll as dry w eight a ccu m ­
ulation and lin e a r growth, have to be c o n sid ered .
The f ir s t v isu a l d e fic ie n c y sym p tom s o c cu rr e d on the - N treatm en t
and w e r e so s e v e r e that 42 ppm o f nitrogen had to be added in o rd er to keep
the plant a liv e .

U ltim ate a n a ly sis o f p e tio le s show ed . 51% and that o f stem

rev e a le d . 63% nitrogen, in oth er w ords, m ore nitrogen w as in the stem s
than in the p e tio le s .

T h e se v a lu e s m u st d efin itely be deficien t, inasm uch

a s lite r a tu r e g e n e ra lly a g r e e s that . 80 to 1. 5% N a re e s s e n tia l for norm al
plant grow th and produ ction.

The low v a lu es e x p r e s s e d th e m se lv e s in

s e v e r e d e p r e ssio n o f lin ea r grow th and dry w eight accum ulation a s w e ll
a s in low ph osphorus and p o ta ssiu m , but rath er high calcium v a lu es in the
p e tio le s .

T he in ter a c tio n w ith p o ta ssiu m and calcium becam e m o re p ro -

75.

nounced under high n itrogen con d ition s.

H ere, the r e v e r s e w as prom inent.

It s e e m s , th e r e fo r e , that d e p r e ssio n o f grow th under high nitrogen le v e ls
m ay be due to antagonism on oth er e le m e n ts and an in d irect e ffe ct of nitrogen
rather than a d ir e c t one, a s is the c a se under nitrogen d eficien t conditions.
V a rio u s sym p tom s of p ota ssiu m d eficien cy , a s d e scr ib e d by W ilhelm
(1950) w e re exp ected , h ow ever, after fiv e w eek s of growth, only the purple
sym p tom s appeared.

T h e se v a rio u s p o ta ssiu m d eficien cy sym ptom s w ill be

d is c u s se d in con n ection w ith E xp erim en t II.

P etio le a n a ly sis and stem analy;

s is from p lan ts tr e a ted w ith p otassiu m d eficien t solution show ed . 12% and
. 30% K r e s p e c tiv e ly , v a lu e s w hich a r e far below th o se rep orted for norm al
field con d ition s.

It is g e n e r a lly accep ted that p o ta ssiu m has a regu latin g

effect on plant m eta b o lism , and is involved in m o st b io lo g ica l rea ctio n s,
how ever, its s p e c ific ity has not been e sta b lish e d .

T h erefore, it is not s u r ­

p r isin g that a s e v e r e p o ta ssiu m d eficien cy w ill d e p r e ss dry w eight accu m u­
lation and lin ea r growth.
The high le v e l of p o ta ssiu m had the sa m e effect on growth as the low
le v e ls .

L eaf sym p tom s, w hich w e r e id en tified by p etio le a n a ly sis a s m a g ­

nesium d eficien cy , app eared a fter nine w eek s o f growth.
sium trea tm en ts,

how ever, did not r ea c t a lik e.

The two high p o ta s­

W hile the le a f sym ptom s

w ere the sa m e, KC1 d e p r e sse d grow th far m ore than K2SO4.

Both tr e a t­

m ents had the sa m e m agn esiu m p e tio le content, how ever, the Cl/SO^ ratio
m ight have been of in flu en ce in asm u ch a s w ith ch lo rid e in the solution, 1. 71%

chloride w as found in the p e tio le s.

T his would be expected to give an addi­

tional d ep ressiv e effect on potassium , which would agree with the findings
of D illey (1957a).

The reduced absorption of both m agnesium and potassium

in the solution containing KC1 may account for growth being le s s for this
treatm ent than for the solution containing ^ S O ^ .
Furtherm ore, the boron p etiole content in both treatm ents was r e la ­
tively very high.

G hrtel (1954) observed weak boron toxicity damage in

vinifera type grapes with high phosphorus and potassium lev els, which
could have occu rred in this case, even though no visual sym ptom s w ere
found.

It could, therefore, be concluded that the depression of growth by

high le v e ls of potassium is due to a combination of factors, such as induced
m agnesium deficiency, as w ell a s extrem e boron, sulfate and chloride
le v els.
One w eek after m agnesium deficiency sym ptom s occurred on the high
potassium treatm ents, sim ila r sym ptom s w ere observed on the basal leaves
of m agnesium deficient plants.

The p etio les and stem s of these plants showed

. 06 and . 07% m agnesium , resp ectively, which is low and below the critical
point of about . 2% (Kenworthy, 1956).

Magnesium plays an important role

as a constituent of chlorophyll, which, in turn, is involved in photosynthesis.
Inasmuch as m agnesium is v ery m obile and e a sily translocated from the
basal lea v es to the term in als, it is understandable that there was very little

d ep ression in lin ear growth.

Furtherm ore, nitrogen and phosphorus w ere

sligh tly higher and calcium was tw o-thirds higher than in the p etioles of
the check; potassium w as somewhat low er while the m inor elem ents seem ed
to be little affected.

The somewhat stronger depression of total dry weight

accum ulation m ight a lso be explained on this b asis.
T here w as a large difference, however, in the shoot/root ratio between
- Mg and 5 Mg.

The latter had the la rg est ratio and was a lso m ore depressing

in the total lin ear growth and total dry weight accumulation.

If dry weight a c ­

cum ulations of the vaious plant parts are compared on the b asis of per cent
of the total, the la rg est difference is found in the roots where 5 Mg had 46%
and - Mg had 20%, w hile the check produced 26%. Petiole an alysis showed
that 5 Mg resu lted in a d ep ression of calcium and potassium , but in creased
phosphorus and boron.

Experim ent II w ill deal with these interactions.

Calcium d eficiency sym ptom s w ere found to coincide with . 26% c a l­
cium p etiole content, and agreed with the reports of Hagler and Scott (1953)
and Steilw aag and Knickmann (1955).

Since calcium is part of the c ell com ­

position and very im m obile, the breakdown occurred in the term inal regions.
Very low calcium content in the p etio les was combined with high nitrogen,
phosphorus, potassium and boron, as w ell as in creased magnesium and
m anganese le v e ls.

Since nitrogen, phosphorus and potassium are present

in large quantities and m agnesium is above normal, the m etabolic p ro cess

of the plant can go on except for the fact that the main c e ll wall constituent
is m issin g without which no elongation or accum ulation can occur.

Also,

the development of the shoots from a xillary buds can thus be explained.
Further d iscu ssio n of calcium in relation to growth w ill be found in E x­
perim ent II.
A fter tw elve w eeks of growth, phosphorus deficiency symptoms
appeared on -P plants.

Petiole and stem an alysis showed . 10 and . 11%

phosphorus, resp ectiv ely .

No phosphorus deficiency has as yet been ob­

served in Concord grape growing regions, and a petiole content of . 2%
is accepted as norm al.

Phosphorus is the source of energy in the m eta­

bolic p r o c e ss and any d eficiency should autom atically slow down the growth
p ro c ess.

It se e m s that deficient le v e ls of phosphorus are coupled with

v ery low nitrogen va lu es.

A d epression in potassium but rather high lev els

of calcium occurred at the sam e tim e.
High le v e ls of phosphorus in the solution produced nearly as much
linear growth as the check, but total dry weight accumulation was depressed.
High le v e ls of nitrogen and boron as w ell as low le v e ls of calcium w ere
present under this condition.

The difference between total linear growth

and dry weight accum ulation might be explained by the presen ce of high
nitrogen and high boron.

High nitrogen furthers linear growth (Kraus and

Kraybill, 1913) and boron, according to the review of Gauch and Dugger (1954)

79.

g iv es the c e ll e la stic ity in addition to having other functions.

Inasmuch as

elongation is enhanced under this condition, the c e lls may be stretched with
le s s than norm al calcium in the w alls, which, in turn, would produce le s s
dry weight.
The experim ent showed under direct or indirect major elem ent defi­
ciency conditions that, in every instance, the stem com position was at lea st
equal to, if not higher than, the p etiole com position, as shown in Table 3.
The amount of nitrogen in stem s and p etioles of the check treatm ent
was very clo se , 1. 05 and 1. 06% resp ectively.

Based on the com parison

with the deficient treatm ents, it must be concluded tliat the check treatm ent
was below norm al for nitrogen and probably m ore growth could have been
achieved with m ore nitrogen present in the plant.
Minor elem en ts w ere of le s s influence on linear growth and dry weight
accum ulation than the m ajor elem ents.

This general effect may have been

due to a m ore sufficient r e se r v e supply of the m inor elem ents within the
plants.

This r e se r v e supply had to be diluted before growth was affected.
M anganese, boron, and copper deficiency sym ptom s appeared at the

sam e tim e.

Plants with m anganese deficiency symptoms had 18 ppm Mn in

p etio les and 23 ppm in stem s.

Beattie and F orshey (1954) reported that m an­

ganese d eficiency can occur when the July p etiole content drops below 30 ppm.
Maume and Dulac (1952) found very large variations of m anganese content in

80.

TABLE 3
COMPARISON BETWEEN STEM AND PETIOLE CONTENT OF ELEMENTS
DEFICIENT OR APPROACHING DEFICIENCY IN THE PLANTS
Specific E le ­
Treatm ent ment D eficient

Content Per Cent - Dry Weight
P etioles
Stems

V isible D e fic i­
ency Symptoms

-N

N

. 51

. 63

Nitrogen

-P

N

. 62

. 62

Phosphorus

-P

P

. 10

. 11

Phosphorus

-K

K

. 12

.27

Potassium

5N

K

1. 57

1. 87

Unidentified

5 K(S04)

Mg

. 14

. 18

Magnesium

5 K(C1)

Mg

. 14

. 19

Magnesium

-M g

Mg

.0 6

.0 7

Magnesium

- Mn

Mn

. 0018

. 0023

Manganese

-K

Zn

. 0025

. 0033

Potassium base
zinc term inals

plants within the sam e grape variety and cam e to the conclusion that location
and so il are the main ca u ses of variation.

The 323 ppm of m anganese of the

5 Mn treatm ent was below the values found in normal field sam ples.

Inasmuch

as Experim ent III deals with m anganese m ore sp ecifically, the resp ective d is ­
cussion w ill be given later.

The standard solution, however, was definitely

too low in m anganese content.

81.

P etio les of boron deficient plants had a content of 14 ppm, and stem s
of the sam e plants had 11 ppm boron.

According to Scott (1954) the range of

deficiency under field conditions lie s between 10 and 20 ppm, while G^rtel
(1956) felt the transition zone should be between 13 and 15 ppm.

Since boron

deficiency im pairs photosynthesis and translocation of sugars from leaves
due to disarrangem ent of the phloem (Reed, 1947), a growth depression and
breakdown of term inal portions m ust be expected.
Even though copper deficiency sym ptom s w ere present, the petiole
an alysis did not show it.

Both - Cu and 5 Cu produced m ore total dry weight

than the check; however, in total linear growth both w ere below the check.
Comparison of p etiole and stem an alysis showed that in m ost treatm ents the
stem copper content was higher than the p etiole content.

Based on the com ­

parison between p etio les and stem s for m ajor elem ents, this would indicate
that copper was deficient in m ost treatm ents.

This w ill be d iscussed further

in Experim ent II.
D eficient plants had m ore zinc in their p etioles than plants fed with
5Zn solution, but deficiency sym ptom s w ere definitely present.

The only

treatm ent resu ltin g in le s s zinc in the p etioles than in the stem s was potassium .
Zinc has an e sse n tia l function in the production of growth regulating substance
which takes place in the m eristem atic regions of the plant.

The p ossib ility

ex ists that sym ptom s ob served on the term inal portions of the - K treatm ent

82.

plants w ere due to zinc shortage because petiole an alysis showed 25 ppm
zinc in the p etio les, which w as the low est concentration found in any trea t­
ment.

Furtherm ore, the zinc content of stem s from the - K treatm ent was

33 ppm.

Sim ilar sym ptom s observed on the term inals of the 5N treatm ent

w ere a lso b elieved to be zinc deficiency, since high le v e ls of nitrogen brought
a se v ere d ep ression of potassium , which, in turn, seem ed to be related to
the zinc content of the p etio les.
cate a deficient zinc lev el.

The p etiole analysis, however, did not indi­

On the other hand, since the p etioles of the

whole plant w ere analyzed together, and, therefore, the values would be an
indication of their average zinc content, they would not show the deficient
level of the term inal portion.
Both - Mo and 5 Mo had som e dep ressin g effect on total linear growth
but 5 Mo produced m ost dry weight, while - Mo produced slightly le s s than
the check.

Petiole an a ly sis showed that both w ere very high in calcium; - Mo

was low in boron, while 5 Mo was slightly above the check.

A combination

of high calcium , a better utilization of nitrate nitrogen for which molybdenum
is essen tia l (Bergman and Kenworthy, 1956) and sufficien nitrogen, phos­
phorus, potassium , m agnesium , and boron w ere probably responsible for
the high dry w eight accum ulation.

The - Mo plants w ere low in molybdenum

and boron in com bination with high calcium and low er potassium concen­
trations.

83.

Total dry weight accum ulation was little affected by iron deficient
nutrient solution, but total lin ear growth was depressed.
d ep ressed both m easu rem en ts.

High le v e ls of iron

In m ost treatm ents iron was higher in the

stem s than in the p e tio les, which would suggest that iron was low or deficient
in the check solution.
of influence.

However, iron source a.nd poor translocation might be

Both low and high iron treatm ents w ere about equal in per cent

distribution of the plant parts and in the shoot/root ratios.

There was, how­

ever, a v ery distinct difference in the appearance of the root system with
high iron plants having v ery poor, very brittle and many dead roots, while
- Fe had a poor root system with black roots, but brittlen ess and dead ones
w ere m issin g .

Both treatm ents produced high calcium lev els in the p etioles

and the potassium value with 5 Fe was 1% lower than the check.

This p otas­

sium d ep ression se e m s to match the high iron value under potassium d efi­
cient conditions.
The total dry weight accum ulation and total linear growth are p resen t­
ed in form of cu rves in Figure 9.

T h ese cu rves are based on the law of dim ­

inishing return with the treatm ents giving highest resu lts in each case at the
apex and the other treatm ents arranged in depressing order towards the base
on each side.

As shown in Figure 9, the relative effect of a specific elem ent

upon dry weight accum ulation or linear growth depends on the sp ecificity of
the elem ent in the m etabolic p r o c e ss and its interaction with other elem en ts.

F ig u r e 9

Top

T otal dry w eight accu m u lation of each of the
25 trea tm en ts arran ged on a cu rve with the
treatm en t producing the la r g e s t amount of dry
weight (5 Mo treatm en t) in the cen ter on the
h ighest point. Low le v e l trea tm en ts w ere a r ­
ranged on the left sid e in d e c r e a sin g order of
d ep ressio n from the b ase lin e tow ards the high­
e st point on the cu rve, and the high lev el treat­
m en ts from th is point tow ards the right in in­
c r e a s in g o rd er of d ep re ssio n . The vertica l
d istan ce from the b ase lin e to any treatm ent
is d irectly p rop ortional to its total dry weight
accum ulation.

Bottom

Total lin ear growth for each of the 25 trea t­
m en ts arranged on a cu rve with the treatm ent
producing m o st total lin ea r growth (check
treatm ent) in the cen ter on the h ighest point.
Low le v e l trea tm en ts w e r e arranged on the
left sid e in d e c r e a sin g o rd er of d ep ression
from the base lin e tow ards the high est point
on the cu rve, and high le v e l trea tm en ts from
th is point tow ards the right in in crea sin g o r ­
der of d e p r e ssio n . The v e r tic a l distance from
the b ase line to any trea tm en t is d irectly pro­
p ortional to its total lin ea r growth.

TOTAL DRY WEIGHT ACCUM ULATION -(G R A M S )

LOW LEVELS

5 Mo

HIGH LEVELS

-Fe
-Cu

CHECK

^ _5K (S 0J
-Co

5P

..5 0

-N
-K -

Leaf symptoms present

TOTAL LENGTH GROWTH-(CM)
LOW LEVELS

CHECK

HIGH LEVELS
■5P

5B
>^5C u

5K (S 04),

5Mn 5Mo

5Fe
*\-5N
5 Co (S04)
f5K(CI)
\5Ca(CI)
5 Mg

♦Leaf symptoms present

85.

The plants used in this experim ent were rooted under normal field condi­
tions and then subjected to sev er e nutritional conditions.

The curves dem on­

strate the value of balanced nutrition (Shear, Crane and M yers, 1946); how­
ever, they do not show the relative effect of an elem ent, should the deficiency
continue to ex ist.

In ev ery ca se this would probably drop the values to zero

with the tim e required depending on the elem ent in question.
Increasing le v e ls of nitrogen, phosphorus, potassium , calcium , m ag­
nesium, m anganese, and boron in the nitrient solution brought an in crease
of each sp ecific elem ent in the p etioles and in stem s.

Iron, copper and zinc

did not follow this tendency. Scharrerand Jung (1956) found a sim ilar situa­
tion in an experim ent with m inor elem en ts on corn and field beans.

With

corn, only iron was irregular, while copper and zinc increased with in crea s­
ing le v els in the nutrient solution.
behaved irregu larly.

In field beans, however, all three elem ents

EXPERIMENT II
E FF E C T S OF
K

RATIOS ON GROWTH, PETIOLE AND STEM
COMPOSITION

In E xperim ent I potassium deficiency appeared on lea v es in the form
of a purple colorin g.

V arious visu al sym ptom s have been reported for pot-

assum d eficiency under field conditions (Wilhelm, 1950).

It has been sug­

gested that th ese various patterns of potassium deficiency may be related
A

to the p resen ce of various le v e ls of calcium and m agnesium .

Experim ent II

was set up to test this p o ssib ility .

EXPERIMENTAL PROCEDURE
The in ter-relation sh ip of calcium , m agnesium and potassium and its
effect upon Concord grape vines was tested with four replications of 18 trea t­
m ents, a s d escribed in Appendix Table 9.

T reatm ents 3 to 18 w ere a fa c­

torial study of four le v e ls of calcium and m agnesium in the nutrient solution
with all other elem en ts being constant.

T reatm ents 1 and 2 contained 117 ppm

potassium , while a ll other treatm ents had only 23 ppm.

Treatm ents 1 and 3

and treatm ents 2 and 13 had the sam e calcium -m agnesium combination, which
made a com parison of the potassium effect p ossib le.
O ne-half strength of the check solution for Experim ent I (Appendix
Table 1) w as used as the b a sis for the experim ent.

The solution was modified,

87.

according to the resu lt of Experim ent I, by doubling the m anganese and in ­
crea sin g the copper content by 50%.
the iron source by Sequestrine.

Also, ferrou s lactate was replaced as

Appendix Table 10 shows the com position

of the 18 solutions and th eir in creasin g Ca+Mg ratios.
K
com position equal to on e-h alf of the check solution.

Treatment 1 had a

T w o-year-old plants w ere stored, pruned, and planted on January 17,
1957, as d escribed in Experim ent I.

After planting they w ere arranged into

four weight groups, and each group was considered as a replication.
average weight per group was 43, 62, 78 and 93 gram s.

The

The plants were

fed for the fir st tim e on February 2.
All but 13 of the plants produced flower clu sters, which were left
on the pLants in order to further the drain of potassium and to see if b erries
have any effect on deficiency sym ptom s.

The flower clu sters were ’’flicked’'

every day to enhance pollination.
When harvested on May 28, 29 and 30, total linear growth and dry
weight accum ulation of roots, stem s, b erries, p etioles and leaf blades were
determ ined.
Experim ent I.

The stem and p etiole tissu e s w ere analyzed as described in
The data w ere sta tistica lly analyzed as a randomized block

experim ent (treatm ents 1 to 18) and as a factorial design (treatm ents 3 to
18).

F ig u r e 10.

Concord grap e le a f injured by A ra m ite sp ray ap plication as
o b serv ed in the green h ou se.

88.

RESULTS

GENERAL OBSERVATIONS
Breaking of buds occu rred very evenly, with the plants of replication
1 (lightest in weight) being a little ahead of the other three replications.

This

gain w as equalized la ter and no difference in rate of growth could be observed.
A spray application of A ram ite (2 -(p-tert-Butylphenoxy) isopropyl-2chloroethyl sulfite) w as used to control a sev ere m ite infestation that developed
%

after four w eeks of growth.

Leaf sym ptom s of injury, as shown in Figure 10,

appeared on m ost le a v es 24 hours later.

A combination of the general con­

ditions of the plants, tem perature and m oisture was suspected as the cause,
because the standard concentration (1 oz. /3 g a l.) that was applied does not
norm ally injure greenhouse plants.

The injured leaves did not drop off dur­

ing the experim ent, although som e of the leaves w ere perm anently injured.
The spray injury made it difficult to fully recognize potassium defi­
ciency sym ptom s on th ese damaged lea v es.

New leaves, unfolding after the

spray damage had occurred, w ere com pletely normal without any effect.
Some lea v es developed the purple castin g as described in Experim ent I.
No so -c a lle d "black leaf" or brown leaves, as have been observed under
field conditions, appeared.

Stem and p etiole an alysis identified the purpling

again as potassium d eficiency.

Some lea v e s on plants receiv in g treatm ents 4 and 6 were very chlorotic
and iron c h lo r o sis w as suspected.

VISUAL APPEARANCE OF ROOTS
T here w as v ery little variation in the general appearance of the root
sy stem s of plants in this experim ent.
with the standard of Experim ent I.

All w ere judged as good in com parison

The only difference between the treatm ents

w as the color, which v aried from dark brown to very light brown.

No brittle

roots w ere found, however, root dry weight accum ulation varied considerably.
Roots of treatm ents 1 and 2 w ere darkest, while those of treatm ent 5
w ere nearly c o lo r le s s .

The roots of treatm ents included in the factorial treat­

m ents appeared as shown in Table 4 (numbers in parentheses are treatm ent
numbers).
TABLE 4
VISUAL APPEARANCE OF ROOTS
Calcium
Concentration - ppm
100
200
301
401

M agnesium Concentration (Ppm)
24
(3)
Dark brown
(7)
Dark brown
(11)
Dark brown
(15)
Dark brQwn

49
(4)
Brown
(8)
Brown
(12)
Brown
(16)
Dark brown

73
(5)
Very light
(9)
Light brown
(13)
Light brown
(] 7)
Dark brown

97
(6)
Brown
(10)
Brown
(14)
Brown
(18)
Light brown

90.

GROWTH

Data pertaining to dry weight of roots, stem s, b erries, p etioles, leaf
blades and total for the plants, together with linear growth, are recorded in
detail in Appendix Table 11.

Dry weight determ inations of the various plant

parts w ere converted to per cent of the total dry weight accumulation for
each treatm ent and sh oot/root ratios w ere calculated.

T hese values are r e ­

corded in Appendix Table 12.
S tatistical a n alysis of the factorial portion of the experim ent showed
no significant d ifferen ces for any of the above mentioned plant parts.

Statis­

tical an a ly sis for the en tire experim ent as a randomized block design of 18
treatm ents showed that the treatm ents having a higher potassium content in
the solution (treatm ents 1 and 2) w ere significantly different from treatm ents
having a low er potassium content in the solution.
Although the

ratio was increased from 5. 3 to 21. 2, there

was no significant difference in the growth of any part of the plant, or the
entire plant.

The variations in the

ratio w ere established by vary­

ing the calcium and m agnesium content of the solution while potassium was
maintained at the sam e lev el (23 ppm).

An in crease in the potassium content

of the solution (ratios 1. 6 and 3. 2) resulted in significantly greater root growth
and greater production of dry m atter for the entire plant.

ELEM ENTA L COMPOSITION OF PETIOLES AND STEMS

D etailed data pertaining to p etiole and stem com position can be found
in Appendix T ables 13 and 14.

Significance of F values, as established in the

sta tistica l a n alysis of the factorial design are reported in Appendix Table 15.
The

ratio in the nutrient solution was of no direct influence

upon nutrient com position of p etio les and stem s for any one element; how­
ever, varied le v e ls of calcium and m agnesium in the solution had a significant
influence upon certain nutrients.

A lso potassium was of influence as shown

by the com parison of treatm ents 1 with 3 and 2 with 13,
Nitrogen:

Large variations in nitrogen content of p etioles and stem s

occurred, even though nitrogen was constant in all solutions.

No statistical

analysis w as made for p etiole nitrogen because of poor petiole growth in som e
treatm ents.
A com parison of treatm ents 1 and 2 (117 ppm K) with treatm ents 3
and 13 (23 ppm K) showed that when calcium and magnesium were at the low
level of supply, and in crea se in the potassium supply resulted in a d ecrease
of nitrogen in stem s and p e tio les.

S tatistical analysis of the factorial design

showed a significant F value for the effect of calcium on nitrogen content of
stem s.

Increasing le v e ls of calcium in the solution produced decreasing

le v els of nitrogen in the stem s, as shown in Table 5.

This influence of

TABLE 5
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON NITRO­
GEN CONTENT (%-DRY WEIGHT) OF STEMS (MEANS OF 4 REPLICATIONS).

Calcium
Cone. (Ppm)

24

M agnesium Concentration (Ppm)
49
73
97

Means for Ca
L evels

100

1 .4 5

1.89

1. 45

1.64

1.68

200

1. 30

1.49

1.4 9

1. 37

1.41

301

1.41

1.32

1.41

1.32

1.37

401

1 .3 3

1.26

1.33

1.27

1. 30

1.41

1.49

1. 46

1.40

M eans for
Mg le v e ls

%

LSD

Calcium le v e ls

.1 2
. 17

5%
1%
NS

Magnesium le v e ls

TABLE 6
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON POTAS­
SIUM CONTENT (%-DRY WEIGHT) OF PETIOLES (MEANS OF 4 REPLICATIONS).
Calcium
Cone. (Ppm)

24

M agnesium Concentration (Ppm)
73
97
49

Means of Ca
L evels

100

.7 6

.8 0

. 66

.7 6

.7 4

200

.6 8

.7 2

.6 4

.4 5

.6 2

301

.5 1

. 46

.5 2

.5 4

.51

401

.7 9

.9 8

. 64

.6 0

.7 5

.6 8

. 74

. 61

.5 9

Means for
Mg le v e ls
LSD

Calcium le v e ls

5%

Magnesium le v e ls

NS

. 19

93.

in creasin g le v e ls of calcium was not observed in the petiole analysis, e x ­
cept where 97 ppm (3x) m agnesium was used.

Here, the nitrogen concentra­

tion of the p e tio le s w as reduced from 3 .1 5 to 2. 43% dry weight as the calcium
supply in the solution was increased.
Potassium :

P etioles of treatm ent 3 (23 ppm K) contained . 76% p otas­

sium com pared to 5. 86% in the p etio les of treatm ent 1 (117 ppm K), or treat­
ment 1 had five tim es m ore potassium in the solution and 7. 7 tim es m ore in
the p etio les than treatm ent 3.

The difference in stem potassium was much

sm aller (. 48% for treatm ent 3 and . 74% for treatm ent 1).

Treatment 2, with

the sam e amount of potassium in the solution as treatm ent 1, but three tim es
m ore calcium and m agnesium , had 3. 21% potassium in the p etioles and . 80%
in the stem s.

T reatm ent 13, on the other hand, which was equal to 2 except

for the low er potassium level, had only . 52% potassium in stem s and p etioles.
S tatistical an a ly sis of the factorial design showed that the calcium
lev els in solution had a significant influence on petiole potassium .

Potassium

d ecreased as the calcium lev el in the solution was increased up to 301 ppm
(3x), but an in crea se to 401 ppm calcium resulted in an in crease of the p otas­
sium content in the p etio les, as shown in Table 6.
The potassium content of stem s was equal to, or higher than, the
potassium content of p etio les for treatm ents that received 301 ppm calcium
(3x) in the solution.

94.

Phosphorus:

T reatm ents 1 and 2, which had 117 ppm potassium in

the solution, had a low er phosphorus content of both p etioles and stem s than
treatm ents 3 and 13, resp ectiv ely (23 ppm potassium ).

Treatment 2 with

three tim es a s much calcium and m agnesium in the solution as treatm ent 1
was significantly higher than treatm ent 1 in phosphorus for both stem s and
p etio les.
The effect of calcium and m agnesium in the solution upon p etioles
(Table 7) and stem s (Table 8) was exp ressed by highly significant F values.
An in crea se in calcium supply resulted in a d ecrease in petiole phosphorus,
*

w hile an in crea se in m agnesium supply caused an in crease in petiole phos­
phorus.

The phosphorus content of the stem s was increased with higher

le v e ls of m agnesium .

A depression in stem phosphorus occurred when the

solution calcium lev el w as raised from 100 to 200 ppm, but further additions
of calcium brought slight in crea ses of phosphorus.
Calcium:

The p etio les of treatm ent 1 had the highest calcium content

(2.40%) of any treatm ent.

Plants of treatm ent 3, which had le s s potassium

in the solution than treatm ent 1, had le s s calcium in the p etioles (. 64%) even
though the calcium supply was equal.

Furtherm ore, treatm ent 2, with the

sam e potassium lev el as treatm ent 1 but three tim es m ore calcium and m ag­
nesium , was low er in petiole calcium (1. 64%) than treatment 1.

Treatment

13 resulted in le s s petiole calcium than treatm ent 2 with equal calcium supply
and le s s potassium supply.

TABLE 7
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON PHOS­
PHORUS CONTENT (%-DRY WEIGH"0 OF PETIOLES (MEANS OF 4 REPLICATIONS).

Calcium
Cone. (Ppm)

24

M agnesium Concentration (Ppm)
49
73
97

Means for Ca
L evels

100

.7 3

. 74

.7 5

.7 7

.7 5

200

.5 0

.6 2

.6 6

.71

.6 2

301

. 57

.5 8

.5 8

.5 8

.58

401

. 52

. 56

.5 3

.5 9

. 55

. 58

. 63

.6 3

.6 6

5%
1.56

.0 5
.0 6

M eans for
Mg le v e ls
LSD

Calcium le v e ls and )
Magnesium le v e ls )

TABLE 8
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON PHOS­
PHOROUS CONTENT (%-DRY WEIGHT) OF STEMS (MEANS OF 4 REPLICATIONS)
Calcium
Cone. (Ppm)

M agnesium Concentration (Ppm)
73
97
24
49

Means for Ca
L evels

100

.2 6

35

.3 0

.3 5

. 31

200

.2 9

27

.2 9

.2 7

.27

301

.2 7

26

.2 9

.2 9

.28

401

.2 9

29

.2 8

.2 9

.2 9

.2 7

30

.2 9

.3 0

Means for
Mg le v e ls
LSD

Calcium and M agnesium ■556
le v e ls
1%

.0 2
.0 3

96.

The calcium lev e l of the solution had a highly significant effect on
stem and p etiole calcium .

The m agnesium level of the solution had a sig n i­

ficant influence on the calcium content of stem s but not on p etioles.

In­

crea sin g calcium le v e ls in the nutrient solution resulted in an increase in
calcium of p etio les (Table 9) and stem s (Table 10).

An in crease in magnesium

in the solution resu lted in a d ecrease of the calcium content in stem s.
M agnesium :

The highest m agnesium content for p etioles was found

in treatm ent 10 (200 ppm Ca, 97 ppm Mg, and 23 ppm K), and the lowest in
treatm ent 1 (100 ppm Ca, 24 ppm Mg, and 117 ppm K).

Treatm ent 2, equal

in potassium content, but with three tim es m ore calcium and magnesium in
the solution than treatm ent 1, had nearly twice as much magnesium in the
p etioles than treatm ent 1.

Treatm ent 13, equal to 2 except for lower potas­

sium, had sligh tly m ore m agnesium in the p etioles than 2.

The stem analysis

followed the sam e trend.
In the factorial experim ent, the m agnesium and calcium lev els had a
highly significant effect on the m agnesium concentration in the p etioles, but
not in the stem s (Table 11).

Increasing lev els of magnesium in the solution,

resulted in an in crea se in petiole m agnesium .

An in crease in the calcium

content of the solution from 100 to 200 ppm resulted in an in crease of the
magnesium content of p etio les.

Increasing the calcium content of the solution

to 301 and 401 ppm resu lted in values for petiole magnesium below that found
for the ones with 200 ppm of calcium in the solution.

TABLE 9
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON CALCIUM
CONTENT (%-DRY WEIGHT) OF PETIOLES (MEANS OF 4 REPLICATIONS).

Calcium
Cone. (Ppm)

M agnesium Concentration (Ppm)
24
49
73
97

Means for Ca
L evels

100

.6 4

.4 5

.4 4

.4 4

.4 9

200

.9 5

.8 5

.8 8

.8 3

.8 7

301

1.0 5

.8 5

.6 2

.71

.8 0

401

1. 05

1.05

.8 4

.7 8

. 93

.9 2

.8 0

.6 9

.68

Means for
Mg le v e ls
LSD

Calcium and m agnesium
le v e ls

5%
1%

. 12
.1 6

TABLE 10
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON CALCIUM
CONTENT (%-DRY WEIGHT) OF STEMS (MEANS OF 4 REPLICATIONS).
Calcium
Cone. (Ppm)

24

M agnesium Concentration (Ppm)
97
73
49
.4 2

Means for Ca
L evels
.41

.4 3

200

. 57

.4 9

. 44

301

.5 5

.5 5

.48

.5 2

. 53

401

.6 4

.6 8

. 59

. 64

.6 0

.5 3

. 54

.4 9

.5 0

Means for
Mg le v e ls
LSD

Calcium and m agnesium
le v e ls

5%
1%

.0 5
.0 7

4s.
00

. 42

•

. 39

4^
to

100

TABLE 11
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON MAGNE­
SIUM CONTENT (%-DRY WEIGHT) OF PETIOLES (MEANS OF 4 REPLICATIONS).

Calcium
Cone. (Ppm)

M agnesium Concentration (Ppm)
24
49
73
97

Means for Ca
Levels

100

.7 4

.8 8

.88

1.2 9

.9 5

200

.7 8

1.10

1.24

1. 71

1.20

301

.8 8

1.29

1. 13

1.25

1. 14

401

.6 9

.8 4

1.08

1. 15

.9 4

. 77

1.03

1.08

1.35

Means for
Mg le v e ls
LSD

Calcium and m agnesium
le v e ls

5%
1%

'

.18
.2 4

TABLE 12
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON MANGA­
NESE CONTENT (PPM-DRY WEIGHT) OF STEMS (MEANS OF 4 REPLICATIONS).
Calcium
Cone. (Ppm)

M agnesium Concentration (Ppm)
97
73
49
24

Means for Ca
L evels

100 .

128

210

169

236

186

200

111

208

165

186

167

301

172

152

139

189

163

401

142

132

111

132

129

138

175

186

146

Means for
Mg le v e ls
LSD

Calcium and m agnesium
le v e ls

5%

1%

27
36

99.

M anganese:

Although the m anganese content of the p etioles varied

between 371 and 890 ppm, the an alysis of variance showed no significant dif­
feren ces between the treatm ents when analyzed as a randomized block design.
However, the a n alysis of the factorial treatm ents showed that calcium
and m agnesium had a significant influence on the m anganese content of stem s
(Table 12).

Increasing le v e ls of calcium in the solution significantly depressed

the m anganese content of stem s.

The fir st three lev els of magnesium in the

solution enhanced the m anganese content in the stem s, but the fourth level de­
p r essed the m anganese content below that found for the 73 and 97 ppm (2x and
3x) le v els.
Iron:

Petiole and stem content of iron was low est in treatm ents 1

and 2, which contained 117 ppm of potassium .

The highest level of potassium

in the solution resu lted in significantly le s s iron in the stem s, regard less of
the lev els of calcium and m agnesium , but had no effect when the calcium and
magnesium was in creased .
The influence of calcium on iron in the stem s was also significant,
with 100 ppm (lx) calcium having the highest iron content and 200 ppm (2x)
calcium having the low est iron content.

The iron content of stem s increased,

however, above that found for 200 ppm, when 300 ppm (3x) and 400 ppm (4x)
calcium w ere added (Table 14).
The calcium -m agnesium interaction was highly significant for iron

TABLE 13
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON IRON
CONTENT (PPM-DRY WEIGHT) OF PETIOLES (MEANS OF 4 REPLICATIONS).

Calcium
Cone. (Ppm)

24

Magnesium Concentration (Ppm)
49
73
97

Means for Ca
L evels

100

101

107

94

83

96

200

75

108

106

103

98

301

124

107

72

85

97

401

86

90

95

98

92

96

103

92

92

NS
5%
1%

30
40

M eans for
Mg le v e ls
LSD

Calcium and m agnesium
le v e ls
Calcium -■magnesium interaction

TABLE 14
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON IRON
CONTENT (PPM-DRY WEIGHT) OF STEMS (MEANS OF 4 REPLICATIONS).
Calcium
Cone. (Ppm)

Magnesium Concentration (Ppm)
97
49
73
24

Means for Ca
L evels

100

105

131

104

131

118

200

85

89

97

85

89

301

86

93

91

104

93

401

99

98

94

101

98

94

103

97

105

M eans for
Mg le v e ls
LSD

Calcium le v e ls
M agnesium le v e ls

5S6
1%
NS

20
26

101.

in p e tio le s (T able 13).

With 100 ppm (lx ) calcium in the solution there w as

no sig n ifica n t in flu en ce of the m agnesium le v e ls upon the iron content of the
p e tio le s .

H ow ever, the lo w e st iron content w as found in com bination with

73 ppm (3x) m agn esiu m .

With 200 ppm (2x) calcium in the solution the u se of

24 ppm (lx ) m agn esiu m r esu lted in iron being sig n ifica n tly below that found
for e ith e r 49 or 73 ppm (2x and 3x) m agn esiu m .

With 301 ppm (3x) calcium

in the solu tion and 73 ppm (3K) m agn esium , the lo w est iron content of p e tio le s
resu lted , w h ile w ith 24 ppm (lx ) m agnesium the h igh est content o ccu rred .
The va rio u s m agn esium le v e ls w ere of no influence upon the iron p etio le
content when com bined with 401 ppm (4x) calciu m .
Boron:

A d e c r e a s e in p ota ssiu m from 117 ppm (treatm ent 1) to 23

ppm (treatm en t 3) r e s u lte d in an in c r r a se in the boron content o f the p e tio le s .
However, w ith higher le v e ls of calcium and m agnesium reduced potassium
le v e ls did not a ffect the am ount of boron in the p e tio le s.
S ta tistic a l a n a ly sis of the fa cto ria l d esign showed that the le v e ls of
calcium and the ca lciu m -m a g n esiu m interaction had a sign ifican t influence
upon the boron content of the p e tio le s (T able 15) and stem s (Table 16).

The

le v el of m agnesium on the oth er hand, sig n ifica n tly affected boron only in
the ste m s.
The in teraction of m agnesium with calcium showed that the influence
of calcium upon the boron content of the p e tio le s w as reduced w ith higher

102.

TABLE 15
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON BORON
CONTENT (PPM-DRY WEIGHT) OF PETIOLES (MEANS OF 4 REPLICATIONS).

Calcium
Cone. (Ppm)

24

Magnesium Concentration (Ppm)
49
73
97

Means for Ca
L evels

100

92

49

58

49

62

200

29

32

30

31

31

301

12

24

32

34

25

401

31

42

27

32

33

41

37

37

37

M eans for
Mg le v e ls
LSD

Calcium le v e ls

5%

6
8

i%

M agnesium le v e ls NS
C alcium -m agnesium interaction

*

12
16

5%
1%

TABLE 16
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON BORON
CONTENT (PPM-DRY WEIGHT) OF STEMS (MEANS OF 4 REPLICATIONS).
Calcium
Cone. (Ppm)

24

M agnesium Concentration (Ppm)
97
73
49

Means for Ca
L evels

100

11

25

25

36

24

200

13

11

21

17

16

301

10

14

19

20

16

401

16

20

12

13

15

12

17

19

22

Means for
Mg le v e ls
LSD

Calcium and m agnesium
5%
le v e ls
1%
C alcium -m agnesium interaction 5%
1%

3
4
6
9

le v e ls of m agnesium .

The low le v e l of calcium resulted in a greater amount

of boron in the p etio les than found for the other lev els of calcium regard less
of m angesium supply.

C onversely the influence of magnesium upon boron

content of the stem s was reduced with higher lev els of calcium .

An in crease

in the m agnesium supply resu lted in an in crease in stem boron only when in
combination with 100 ppm (lx ) of calcium .
Copper:

The use of 117 ppm of potassium resulted in a significantly

higher value for copper in the p etioles than found with 23 ppm of potassium
when the le v el of calcium and m agnesium in the solution was low (lx).
Increasing the le v el of calcium or m agnesium in the nutrient solution
had a significant d ep ressin g effect on copper in the p etioles (Table 17).

The

influence of calcium w as a sso cia ted m ainly with the low level (lx).
Furtherm ore, the F value for the interaction calcium -m agnesium
of the factorial design indicated that a significant influence was exerted on
the copper content of both p etio les and stem s.

The use of 24 and 49 ppm

(lx and 2x) m agnesium d ep ressed copper in p etioles significantly more
when combined with 200, 301 and 401 ppm (2x, 3x and 4x) of calcium than
when combined with 100 ppm (lx ) calcium in solution.

The combination of

73 ppm (3x) m agnesium with 200 and 401 ppm (2x and 4x) of calcium in the
solution d ep ressed copper m ore than when combined with 100 or 301 ppm
(lx and 3x) in the solution.

A ll four calcium le v e ls in combination with

TABLE 17
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON COPPER
CONTENT (PPM-DRY WEIGHT) OF PETIOLES (MEANS OF 4 REPLICATIONS).

Calcium
Cone. (Ppm)

24

M agnesium Concentration (Ppm)
73
97
49

Means for Ca
L evels

100

37

29

27

25

29

200

23

23

20

23

22

301

23

24

27

23

24

401

25

26

21

24

24

27

25

23

23

Means for
Mg lev e ls
LSD

Calcium and m agnesium
5%
le v e ls
1%
C alcium -m agnesium interaction 5%
1%

2
3
4
6

TABLE 18
INFLUENCE OF VARIABLE CALCIUM AND MAGNESIUM LEVELS ON COPPER
CONTENT (PPM-DRY WEIGHT) OF STEMS (MEANS OF 4 REPLICATIONS).
Calcium
Cone. (Ppm)

M agnesium Concentration (Ppm)
97
73
24
49

Means for Ca
L evels

100

20

21

23

20

21

200

21

20

21

17

19

301

16

19

23

23

20

401

19

22

18

20

20

18

20

21

20

Means for
Mg le v e ls
LSD

Calcium and m agnesium lev els
C alcium -m agnesium interaction

NS
5%
1%

3
4

97 ppm (4x) m agnesium resu lted in alm ost the sam e copper values for
p etio les.
In stem a n a ly sis it w as found that the lx lev el of magnesium when
com bined with the lx lev el of calcium in the solution depressed copper sign i­
ficantly m ore than when combined with the other three calcium lev els (Table
18).

With 49 and 73 ppm (2x and 3x) magnesium there was a significant dif­

ference between the 3x and 4x le v e ls of calcium .

97 ppm (4x) magnesium

combined with 200 ppm (2x) calcium in the solution d epressed copper uptake
m ore than when com bined with the other three calcium lev els.
Zinc:

The treatm ents (1 and 2) that contained 117 ppm potassium

resulted in significantly m ore zinc in the p etio les than the corresponding
treatm ents (3 and 13) that contained only 23 ppm potassium in the solution.
An in crea se in the supply of calcium and m agnesium significantly reduced
the amount of p etiole zin c a sso ciated with the use of 117 ppm of potassium .
No significant influence of calcium or magnesium supply upon zinc
content of either p etio les or stem s was found in the factorial experim ent.

106.

DISCUSSION

A v a ria tio n in the app earance of p otassiu m sym ptom s o ccu rs not only
on la b ru sca type grape p lan ts, but a lso on v in ifera v a r ie tie s .
H arding (1957) o b se rv e d only purple sym ptom s in an experim en t on
C oncord g ra p es w ith nutrient culture.

Stellw aag (1955), in nutrient culture

ex p erim en ts with R ieslin g , Sylvaner, and Burgunder v in es, was able to
produce brown sym p tom s by u sin g Van der Crone, H e llrieg el, and
M erk en sch la g er so lu tio n s.

He w as, how ever, unable to obtain them with

Hoagland solution b ecau se iron c h lo r o sis appeared and m asked the sym ptom s
of p otassiu m d e ficien cy .

The statem ent of G&rtel (1955) that with potassium

d eficien cy sym ptom s the ratio of K^OiMg in le a v e s is below 1, held true
for p e tio le s in th is ex p erim en t.
Knotte and R itsch l (1931) gave a d escrip tio n of nutrient culture r e ­
su lts obtained under p otassiu m d eficien t conditions.

They m entioned that

" g rea sy looking spots" appeared fir s t on the le a f ep id erm is between the
v ein s.
off.

T h ese sp ots la te r turned yellow and then brown, with the tissu e dying

P relim in a ry ex p e rim en ts by the author (Bergman and Kenworthy, 1956)

showed s im ila r sym p tom s under com bined molybdenum and potassium d e fi­
cien t conditions.

Inasm uch a s in 1931 not m uch w as known of the e ss e n tia lity

of m olybdenum for plant growth, it m ight w ell be worth w hile to further in ­
v e stig a te th ese p o s s ib le con n ection s.

107.

The com position of the nutrient solutions in this experim ent was
designed to study various Ca.+Mg ratios in r e iation to the various potassium
K
deficiency sym ptom s d escribed in literature.

Since in Experim ent I the

potassium content of the p etio les was far above the usual field potassium
content, it w as felt that l / 2 x m odified Hoagland solution would be a ju sti­
fied b a sis.

However, it becam e evident from the stem and p etiole analysis

at the end of the experim ent that 23 ppm potassium in the solutions for trea t­
m ents 3 to 18 was too low in com parison with the calcium and magnesium
le v els and that potassium m ay have been the lim iting factor in all treatm ents.
On the other hand, if higher le v e ls of potassium would have been used, the
calcium and m agnesium should have been increased to provide the desired
ratios and potassium may still have been the lim iting factor.
The higher Ca+Mg ratios obtained by in creasin g the calcium and
IC
m agnesium supply had no direct influence on the stem or petiole com position
for any sp ecific elem ent.

Increasing ratios of treatm ents 1, 2 and 3 (ratios

1. 6 to 5. 3) resu lted in a d ecrea se in potassium , calcium and zinc and an
in crease in nitrogen, phosphorus, boron and copper content of the p etioles.
Furtherm ore, there w as an in crease in total dry weight and an in crease in
total linear growth and in the shoot/root ratios.

Since treatm ents 1 and 2

had the sam e potassium content in the solution, with 2 having higher calcium
and m agnesium le v e ls, and treatm ent 3 having lower potassium content than 1,

10 8 .

the e ffe c t se e m e d to be due to p o ta ssiu m rath er than the r a tio s.

O therw ise,

trea tm en ts 3 to 18 should have shown s im ila r resp o n se .
S ta tistic a l a n a ly s is of the fa cto ria l d esign (treatm en ts 3 to 18), in
w hich only the calciu m and m agnesium le v e l of the solution w as varied , gave
som e in d ication s of how the v ariou s e lem en ts in p e tio le s and ste m s w ere
affected by th e se two e le m e n ts or th eir in teraction (Appendix T able 15).

In­

c r e a sin g le v e ls of calciu m in the solution resu lted in a sign ifican t in c r ea se
in calcium and a d e c r e a s e in phosphorus, p otassiu m , boron and copper in
p e tio le s.

In crea sin g le v e ls of m agnesium brought a sign ifican t in c r e a se of

phosphorus and m agn esiu m , but a d e c r e a se in calcium and copper.
ste m s follow ed the sam e trend.

The

The e ffe ct of calcium and m agnesium in

the solution upon th ese e lem en ts in p e tio le s or ste m s, ag reed w ith the r e ­
su lts of E xp erim en t I.
It is in te r e stin g to note that when the potassiu m le v el in the solution
w as red uced from 117 ppm to 23 ppm, th ere w as a slight in c r e a se in the m a g ­
nesium content of the p e tio le s and the calcium content of the p e tio le s de­
c r e a se d from 2. 40 to . 64% (treatm en ts 1 and 3) w hile with a higher le v e l of
calcium in the solution a fu rther d e c r e a se from 1. 64 to 1. 13% (treatm ents
2 and 13) w as o b serv ed .

T h is could be in terp reted that a certa in le v el of

p otassium in the su b stra te m ay be n e c e s s a r y for an adequate absorption of
calciu m .

109.

In the chapter on d e fic ie n c y and to x icity sym ptom s of le a v es, it w as
stated that tr e a tm e n ts 4 and 6 w ere ch lo ro tic and iron c h lo r o sis w as expected
to be the c a u se.

Both trea tm en ts, how ever, had the h igh est iron content (131

ppm) of the s te m s.

Gouny and M azoyer (1953) felt that there are two d iffe r ­

ent kinds of iron c h lo r o sis: one, w here the iron is tied up in the so il due to
high pH, and the oth er due to m eta b o lic action s w ithin the plant.

In the latter

c a se , v e r y low le v e ls or v e r y high le v e ls of potassiu m m ight be resp o n sib le.
Lindner and H arley (1944) found v ery high p otassiu m concen trations in le a v es
of Bartlett p e a r tr e e s show ing iron c h lo r o sis.

The data of E xperim ent II

show that the above m entioned fir s t rea so n could not be resp o n sib le for the
o b se rv e d c h lo r o s is b eca u se the pH of the solution w as 5. 0 and, furtherm ore,
actu ally m ore iron w as found in the ste m s of the ch lo ro tic plants than in the
other ste m s.
H ow ever, one strik in g factor is the high le v e l of iron found in the
ste m s, a s com pared to the low er le v e ls found in p e tio le s.

T his phenomena

could not be a s so c ia te d with any sp e c ific ratio of ca lciu m /p o ta ssiu m , m agcalcium + m agnesium
n esiu m /p o ta ssiu m , ca lciu m /m a g n e siu m or ----------;------------- .
'^
potassium

^
E xp eri-

m ent I produced the h igh est iron s te m value under p otassiu m d eficient condi­
tion s (100 ppm v e r s u s 80 ppm in p e tio le s), w h ile the high calcium treatm ent
had slig h tly m ore iron in the p e tio le s than in the stem s (50 ppm v e r su s 45 ppm).
F u rth erm ore, in th is exp erim en t, the h igh est phosphorus p etio le and stem
contents co in cid ed w ith the h igh est iron stem contents.

110.

Based on th ese data, it is suggested that under low or potassium defi­
cient conditions the m etabolic p ro c e ss in the plant is slowed down.

Since this

slow er p r o c e ss needs le s s energy, derived from the organic phosphate which,
in turn, is p resen t within the plant in higher concentration, any e x c e s s would
com bine with the iron and form an insoluble iron-phosphate com plex which
would prevent any further iron translocation to the p etioles.
To the author's knowledge nobody has been able to explain the various
potassium d eficiency sym ptom s on grape vin es from a nutritional viewpoint.
The r e su lts of this experim ent suggest that this problem may not be solved
on the b a sis of m ineral nutrition.

The occurrence of necrotic tissu e s in

the event of potassium deficiency may be considered as the normal symptom.
An investigation of the various lea f colors associated with potassium defi­
ciency may b est be approached on a biochem ical b asis to estab lish what
pigm ents are involved and to study the quantities present, as w ell as to
investigate the origin of the pigm ents.

An approach sim ilar to that employed

by Burghardt (1956) could be used as b asis for tests in this respect.

111.

EXPERIMENT III

EFFEC T OF THREE LEVELS OF MANGANESE ON GROWTH, PETIOLE AND
STEM COMPOSITION
The p etiole m anganese content of the check treatm ent in Experiment
I w as very low, com pared to the average of field sam ples collected in late
July.

It w as also found that the m anganese content of p etioles from plants

grown under field conditions in creased throughout the season (Bergman and
Kenworthy, 1957), and reached a value as high as 2700 ppm in late fall with­
out any sp ecific leaf sym ptom s.

T herefore, Experiment III was designed

to find out how high the m anganese content in the solution had to be in order
to reproduce the values for m anganese found in average field sam ples and
to observe the effects of very high le v e ls of m anganese in the solution upon
the plants and absorption of nutrients.

EXPERIMENTAL PROCEDURE
This experim ent w as conducted sim ultaneously with Experiment II.
The experim ental m ethods w ere the sam e as for Experiment II.

Further­

m ore, the check of Experim ent II ( l/2 x m odified Hoagland solution with 1. 0
ppm (2x) m anganese) serv ed a lso as a check for this experim ent (Appendix
Table 9).
The other treatm ents had 12. 5 and 25 ppm (25x and 50x Hoagland)
m anganese in the l / 2 x Hoagland solution.

112.

RESULTS
GENERAL OBSERVATIONS
S in ce the ex p erim en t w as c a rr ie d out at the sam e tim e and with plants
lik e th ose of E x p erim en t II, the gen era l behavior o f the plants, including spray
injury, w as the sa m e.

H ere too, the flow er c lu s te r s w ere left on the plants

and the b e r r ie s w e re allow ed to develop.

On warm , sunny days, the plants

supplied w ith 12. 5 and 25 ppm m an ganese seem ed to need m ore sulution b e­
cau se w iltin g o c cu rr e d m ore quickly.

T his may have been a s so c ia te d with

the rath er fin e textu re of the young le a v e s.

Plants r e c e iv in g the high le v e ls

of m an gan ese grew v ery w ell, but the b e r r ie s did not seem to ripen evenly
on each c lu ste r , w hile they ripened w ell on the plants of the ch eck treatm en ts.

VISUAL SYMPTOMS
D uring 15 w eek s, the high m anganese plants grew norm ally like the
check p lan ts.

At th is tim e the youngest le a v e s of the high (25 ppm) m anganese

treatm en t began to show in tervein a l c h lo r o sis with the v ein s staying green
(F igu re 11).

A fter a w eek, the green veination disappeared too, and the

e n tire le a f took on a g reen ish -b ro w n co lo r w ith darker brown ed g es not w ider
than the depth of the dentation.
c o lo r and did not turn brow nish.

The old er le a v e s kept th eir norm al green
L eaf enlargem en t w as sligh tly ham pered.

No sim ila r sym ptom w as o b serv ed throughout th ese in v estig a tio n s.

The

roots w ere judged a s good, and actu a lly nothing sp e c ific w as ob serv ed on either
stem or r o o ts.

113.

F igu re 11.

Symptom on grape le a v e s o b serv ed when plants w ere
supplied w ith the high lev el (25 ppm) of m anganese in
the nutrient solution, after 15 w eek s of growth.

MANGANESE

LEVEL IN SOLUTION g M

|

PPM (CHECK)

17 / f * 12.5 PPM
2 5 PPM

GROWTH— CM

600
150
500
400

TOTAL

LENGTH

300
200

I OC

l

3
«
or

I
h- 100
CO

§
§

DRY WEIGHT-SRAM

I

I
12.5 25
SOLUTION CONTENT

F ig u r e 12.

5C*

I
.
I
125 25
SOLUTION CONTENT

I
12.5 25
SOLUTION CONTENT

PETIOLES

1

CO

LEAF BLADES

El] 20

SOLUTION

CONTENT

E ffe ct of th ree le v e ls of m an gan ese in the solu tion on total
lin ea r growth, total dry w eight, p e tio le and le a f blade dry
w eight accu m u lation .

114 .

GROWTH
Data p erta in in g to total lin ea r growth and total dry weight accu m u la­
tion of ro o ts, ste m s, b e r r ie s , p e tio le s and lea f blad es can be found in Appendix
T able 16.

D ry w eight accu m ulation of roots, ste m s and b e r r ie s, p e tio le s and

le a f b lad es con verted into p er cent of total dry w eight accum ulation are r e ­
cord ed in Appendix T able 17.

T otal L inear Growth (F igu re 12)
T here w as a sign ifican t in c r e a se in total lin ea r growth from 192 cm
to 658 cm when the m an gan ese content in the solution w as in crea sed from
1. 0 ppm to 12. 5 ppm.

H ow ever, further in c r e a s e s in m anganese brought a

sligh t d e p r e ssio n to 610 cm which, how ever, w as still sign ifican tly larger
than the check .

T otal D ry W eight A ccum ulation (F igure 12)
The total dry w eight accum ulation in c r ea se d with in crea sin g m angan­
e s e content in the solution .

The treatm en t containing 25 ppm m anganese in

the solution produced sig n ifica n tly m ore total dry w eight than the check ( 1. 0
ppm Mn) treatm en t.
dry m atter.

T he 12. 5 ppm treatm ent resu lted in 150. 36 gram s of

T his w as 31 g ra m s m ore than found for the check, and 37 gram s

le s s than for the treatm ent containing 25 ppm m anganese.

D ry W eight A ccu m ulation of R oots, S tem s, B erries, P e tio les and L eaf Blades
S ta tistic a l a n a ly s is for dry w eights of the above m entioned plant p arts
show ed no sig n ifica n t d ifferen ce betw een the trea tm en ts for roots, stem s
and b e r r ie s .

The dry w eight of ste m s, how ever, in c r e a se d with in crea sin g

le v e ls of m an gan ese in the solution .

The 12. 5 ppm treatm ent produced tw ice

as m uch stem dry w eight and the 25 ppm treatm ent three tim es a s much as
the check.
T his sa m e trend held true for the p e tio le s.

H ere both the 12. 5 and

25 ppm trea tm en ts w ere sig n ifica n tly higher in p etio le dry w eight than the
check (F igu re 12).

Although the 25 ppm m anganese le v el produced m ore

p e tio le dry w eight than the 12. 5 ppm m anganese le v el, the differen ce was
not sign ifican t.
D ry w eight accu m ulation of lea f blades in c r ea se d with in crea sin g
le v e ls of m an gan ese in the solution.

The use of 12. 5 ppm of m anganese

r esu lted in sig n ifica n tly g r e a te r production of lea f blades than 1. 0 ppm of
m an gan ese (check).

A lso, the u se of 25 ppm of m anganese in the solution

r esu lted in sig n ifica n tly g r e a te r dry w eight of le a f blades than 12. 5 ppm
(F igu re 12).
A ccum ulation of D ry W eight E x p r essed in Per Cent of T otal Dry Weight by
R oots, Stem s and B er r ies, P e tio le s, and L eaf Blades
C onverting the dry w eight m ea su rem en ts into p er cent of total dry
w eight show ed that ste m s and b e r r ie s , p e tio le s, and lea f blades in crea sed

1500

IOOO

MANGANESE

CONTENT-PPM

DRY WEIGHT

PETIOLES

MANGANESE

F ig u r e 13.

IN

NUTRIENT S O L U T I O N - P P M

E ffe c t o f th r e e l e v e l s o f m a n g a n e se in th e s o lu tio n o f
m a n g a n e se and ir o n in th e p e t io l e s .

116.

w ith in c r e a s in g le v e ls of m an gan ese in the solution, w hile the roots d ecrea sed .
Since shoot grow th in c r ea se d , and root growth d ecrea sed , there w as an in ­
c r e a s e in the s h o o t/r o o t r a tio s a s the m an ganese le v e l w as in crea sed .

The

d iffe re n c e s in the p er cent of total dry w eight for the different plant p arts
w as alw ays g r e a te r for the in c r e a se from the 1. 0 to 12. 5 ppm m anganese
le v e l than for the in c r e a s e from the 12. 5 to 25 ppm m anganese.

ELEM ENTAL COMPOSITION OF PETIOLES AND STEMS

D ata p erta in in g to p e tio le and stem com position a re given in Appendix
T ab les 18 and 19, r e s p e c tiv e ly .

S ta tistic a l a n a ly sis showed a significan t effect

of the trea tm en ts on uptake of iron and m an ganese in p e tio le s without any s ig ­
nificant in flu en ce on the oth er e lem en ts.

The elem en ta l com position of the

ste m s w as not sig n ifica n tly affected .
Although the d iffe r e n c e s w ere not sign ifican t the in c r ea se in m anganese
supply see m e d to d e c r e a s e the p etio le concentration of nitrogen, phosphorus,
p otassiu m , calciu m and m agn esiu m .

Stem a n a ly sis for nitrogen showed a

d e c r e a se , w h ile phosphorus, p otassiu m and calcium showed an in c r ea se
with in c r e a sin g le v e ls of m an gan ese in the solution.
Iron (F ig u re 13) in the p e tio le s w as sign ifican tly in c r ea se d as the
m an ganese content of the solution w as in crea sed .

The in c r e a se in iron of

ste m s a s s o c ia te d w ith in c r e a sin g le v e ls of m anganese in the solution was

h ow ever not sig n ifica n t.

Iron in the p e tio le s of the 25 ppm m anganese tr e a t­

m en t w a s sig n ifica n tly high er than in the 12. 5 ppm treatm ent and in the check
(1. 0 ppm Mn).

In a ll th ree treatm en ts the stem iron content w as low er than

the p e tio le iron content.
M anganese (F igu re 13) in p e tio le s w as in c r ea se d fiv e fold when m an­
g a n ese in the solu tion w as in c r e a se d from 1. 0 to 12. 5 ppm.

T h ere w as about

eight tim e s a s m uch m an gan ese in the p e tio le s from the 25 ppm treatm ent
than in the p e tio le s from the ch eck treatm ent (1. 0 ppm Mn).

The ste m s had

th ree tim e s m ore m an gan ese when the solution m anganese le v el w as r a ise d
from 1. 0 to 12. 5 ppm, and sev en tim e s m ore when in c r ea se d to 25 ppm.
A lso, the p e tio le s contained four tim e s m o re m anganese than the stem s
for the ch eck treatm en t, the d ifferen ce in c r e a se d to sev en fold when m an­
gan ese w as in c r e a s e d to 12. 5 ppm, and to 4. 7 fold w ith 25 ppm m anganese in
the solution .

The valu e of 3067 ppm of m anganese in the p e tio le s from the

25 ppm m an gan ese treatm en t w as com parable to the high v a lu es found under
fie ld conditions in la te su m m er.

H ow ever, the u se of 12. 5 ppm of m anganese

in the solu tion s r e s u lte d in v a lu es fo r p etio le m anganese in e x c e s s of the
v a lu es found for a v e ra g e fie ld sa m p les in m id su m m er.

118 .

DISCUSSION

M anganese in the solution strongly influenced the m anganese co n cen ­
tration in the plant, and see m e d to be taken up in solution to its amount
p r e se n t in the su b str a te .

M aume and DuLac (1952) reported, after in v e s ­

tigatin g v a r io u s grape v a r ie tie s grown in different location s, that the s p e c i­
fic ity of a v a r ie ty or s p e c ie s w as not r esp o n sib le for the m anganese content
of the v a rio u s plant p a r ts, but p r im a r ily the location.

L arge d ifferen ces of

m an gan ese in grape p e tio le s w ere found by Bergman and Kenworthy (1957)
under M ichigan conditions w hich coin cid ed w ith highly variable m anganese
le v e ls in M ichigan s o ils (Lawton, 1957).

T h erefo re, the reaction of grapes

to m an ganese in nutrient cultu re is understandable.

Furtherm ore, the pH

of the nutrient solution u sed in this experim en t w as about equal to the pH of
the so il in w hich grap es a re grown in M ichigan.
It is in te r e stin g to note that apples, containing over 500 ppm m an­
ga n ese in the le a v e s, n orm ally show internal bark n e c r o sis (m ea sle s) on
the ste m s (Kenworthy, 1957) w hile under fie ld conditions the grape vine
with 2800 ppm in the p e tio le s produced no o b serv ed sym ptom s.

The leaf

sym ptom s p r e se n t in E xperim en t III with 3067 ppm m anganese in the p e tio les
m ay have been the beginning of a d eficien cy induced by the high le v e ls of
m an gan ese or an e x p r e s s io n of m an ganese e x c e s s .
In s te m s of the 25 ppm m an ganese treatm ent plants, the phosphorus

119.

w a s h igh er than in the p e tio le s w hich would indicate, if interpreted on the
b a sis of fin din gs in E xp erim en t I, that the plants w ere low or deficient in
phosp horus.

E x tr em ely high m anganese le a f v a lu es w ere found under field

conditions by G&rtel (1956) and w ere a sso c ia te d with phosphorus d eficien cy
sym p tom s.

H ow ever, the sym ptom s o b serv ed on the plants of the 25 ppm

treatm en t w ere not s im ila r to sym ptom s of phosphorus d eficien cy found
in E xp erim en t I.
The p o ta ssiu m content of p e tio le s and ste m s d ecrea sed with in crea sin g
m an ganese le v e ls in the solution.

The data from E xperim ent I suggest that

with high le v e ls of p o ta ssiu m in the solution the m anganese w as d ep ressed
m ore than the p otassiu m w as with high m an ganese le v e ls .
T h ere a r e m any rep o rts in liter a tu r e indicating high le v e ls of heavy
m eta ls in the solu tion w hich induce iron c h lo r o sis (Goodall and G regory, 1947;
DeKock, 1956).

W einstein and Robbins (1955) noted that under high m anganese

and low iron le v e ls true iron d eficien cy is induced.

On the other hand,

Burghardt (1956) w as unable to find any ind ications of iro n /m a n g a n ese antagon­
ism in spinach p lan ts.

He based h is statem ent on r e su lts obtained on the rate

of CO2 a ssim ila tio n and resp ir a tio n as w ell as the amount of chlorophyll and
xanthophyll p r e se n t under low and high iron or m anganese le v e ls.

In E x p eri­

m ent III the iron in p e tio le s in c r e a se d with in crea sin g m anganese in the so lu ­
tion s.

F u rth erm ore, the d ifferen ce betw een stem and p etio le iron in crea sed

with h igh er m an gan ese le v e ls .

In trea tm en ts 4 and 6 of E xperim ent II, the

120.

p e tio le iron content w as far below that in ste m s and the v is ib le sym ptom s
(iron c h lo r o s is ) did not m atch the sym ptom s obtained under high m anganese
le v e ls or sym p tom s found in E xperim en t I.
T he c o r r e la tio n co effic ien t betw een iron and m anganese in p e tio les
w as c a lcu la te d sep a ra tely for trea tm en ts v a ried in iron and m anganese (E x ­
p e r im e n ts I and III), calciu m and m agnesium (E xperim ent II), and for all the
tr e a tm e n ts of E xp erim en t I w here e v er y elem en t w as varied and the follow ing
r e s u lts w e re obtained:
only iron and m an ganese varied

+ . 826

a ll elem en ts v a ried

+ .7 1 8

calciu m and m agn esium v a ried

+ .4 6 1

The fir s t two c o rr e la tio n c o e ffic ie n ts w e re sign ifican t at the 1% le v e l and the
third at the 5% le v e l.

T his s u g g e sts that iron and m anganese are a ctin g ind e­

pendently; oth erw ise, a n egative c o rrela tio n should have been found.
Since the c o rr e la tio n betw een iron and m an ganese in the calcium and
m agn esiu m trea tm en ts is com p a ra tiv ely low, a p o ssib le d irect or indirect
rela tio n could e x is t w ith th ese e le m e n ts.

It, th erefo re, appears that the

v isu a l sym p tom s o b se rv e d with the 25 ppm treatm ent are m anganese to x icity
sym ptom s, b ased on sp ectro g ra p h ic a n a ly sis for total iron in p e tio le s and
ste m s .

Since the le a v e s w ere not analyzed and no com p arison w as made b e ­

tw een "active" and total iron p r e se n t (Sam ish, 1954) it is felt that the sym p ­
to m s o b se rv e d w ere not connected with low le v e ls of total iron.

121 .

SUMMARY

The r e sp o n se of Concord grape v in e s to high and low le v e l s of
e le v e n nutrient e le m e n ts w as studied in the greenh ouse in th ree ex p erim en ts.
In each c a se , the p lan ts w e re grown for 16 w eek s with nutrient solution s and
u sin g quartz sand a s grow ing m edia.
In E xp erim en t I o n e -y e a r -o ld rooted cuttings w ere u sed and the check
trea tm en t w as supplied with standard Hoagland solution No. 1 (Hoagland and
Arnon, 1950).

The high and low le v e ls of elev e n nutrient elem en ts w ere ob-

tain ed by ad ju stin g the com p osition of the standard solution to the sp e cific
le v e ls .

High calciu m and p otassiu m w ere te ste d in com bination with sulfate

and ch lo rid e a s anion s.

T h is provided 25 trea tm en ts which w ere arranged in

a random ized block w ith th ree r e p lic a te s per treatm ent.
In E x p erim en ts II and III tw o -y e a r -o ld rooted cuttings w ere used.
The ch eck solution w as m od ified by doubling the m anganese and in crea sin g
the cop p er content by 50%.

R atios from 1 .6 to 21. 2 of

K

in the nutrient

solution, b ased on l / 2 x m od ified Hoagland solution, w ere used to te s t the
in flu en ce on grow th and the v is ib le e x p r e ssio n of potassium d eficien cy a r ­
ranged in a ran d om ized block, 18 trea tm en ts with four r e p lic a te s per tr e a t­
m ent, w e re the b a s is for E xp erim en t II.

Sixteen of th ese 18 treatm en ts w ere

se t up a s a 4 x 4 fa c to r ia l d esign containing four le v e ls each of calcium and
m agn esiu m , w ith a ll oth er e le m e n ts constant.
w e re v a r ie d in the p o ta ssiu m content.

The other two treatm ents

122.

In E xp erim en t III the effect of th ree le v e ls of m anganese in the nutrient
solu tion w as te s te d w ith th ree trea tm en ts having four r e p lic a te s each.

P e tio les

and s te m s of each plant w ere analyzed for nitrogen, phosphorus, potassium ,
ca lciu m , m agn esiu m , iron, m anganese, boron, copper, zinc, and p a rtia lly
for ch lo r id e .
T otal lin ea r growth, total dry w eight accum ulation, a s w ell as dry
w eight accu m u lation by roots, ste m s, b e r r ie s , p e tio le s, and lea f blades w ere
taken.
Low le v e ls of nu trien ts d e p r e sse d dry w eight accum ulation in the
follow in g d e c r e a s in g order: p o ta ssiu m ,> n itro g e n ,> ca lc iu m > p h o sp h o r u s,
>boron, > m a g n esiu m ,> zinc,> m an ganese> m olybdenu m .

High le v e ls of nutrients

w ere of d e p r e ss in g e ffe c t in the follow ing d e c re a sin g order:

potassium with

c h lo r id e ,> m a g n e siu m ,> c a lc iu m with su lfate,> calcium w ith ch lorid e,> n itrogen ,
> p h o sp h o ru s,> p o ta ssiu m w ith su lfa te,> iro n ,> b o ro n ,> m a n g a n ese.

The high

m olybdenum treatm en t r e su lte d in the g r ea test dry w eight accum ulation.
L inear grow th w as d e p r e sse d by d eficien t le v e ls of nutrients in the
follow in g d e c r e a sin g order: n itro g e n ,> p o ta ssiu m /> p h o sp h o ru s> ca lciu m ,
>boron,>.copper, > zin c,> m olybdenum ,>iron2> m an gan ese,> m agn esiu m .

High

le v e ls of n u trien ts d e p r e sse d in the follow ing d ecrea sin g order: m agnesium ,
> calciu m and p o ta ssiu m with c h lo r id e ,> calcium with su lfa te,> n itro g en ,> iro n ,
> m a n gan ese,> m olyb d en u m ,> z in c ,> p otassiu m with sulfate,> copper,> boron.
The ch eck treatm en t r e su lte d in the g r e a te st lin ea r growth w ith the high
ph osphorus treatm en t c lo s e second.

T re a tm en ts d eficien t in n itrogen and phosphorus or high in m agnesium
and m olybdenum produced the lo w e st sh o o t/r o o t r a tio s.

High le v e ls of p h o s­

ph oru s, n itrogen and low le v e ls of m agnesium resu lted in h igh est sh o o t/ro o t
r a tio s .
Based on dry w eigh t accum ulation, low p otassiu m , high calcium with
ch lo rid e, and high m agn esium produced le a s t p e tio le s p e r c e n ta g e -w ise .
M ost le a f b lad es (per cen t of total dry w eight) w ere cau sed by high nitrogen
le v e ls .
A p o s itiv e rela tio n sh ip betw een the sp e c ific nutrient elem en t in the
solu tion and the p e tio le content w a s o b serv ed for a ll e lem en ts with the e x ­
cep tion of iron and z in c .

P e tio le s see m e d to be the b etter in d icators of

norm al and above norm al le v e ls of nutrition than the ste m s.

The ste m s, on

the oth er hand, w e re b etter in d ica to rs of low and d eficien t le v e ls .

The stem

content w as alw ays equal to, o r higher than, the p etio le content under low
o r d e fic ie n t nutrient con d ition s.
The

K.

r a tio s in the nutrient solution w ere of no d irect influence

on total lin ea r growth, total dry w eight accum ulation, or the appearance of
the p o ta ssiu m d e fic ie n c y sym p tom s.

Indirectly, how ever, potassium , c a l­

cium or m agn esiu m e x e r te d v a r ied in flu en ces.
C h ronological app earance of v is ib le d eficien cy or to x icity sym ptom s
did not c o r r e la te w ith the d e p r e ss iv e elem en ta l e ffe c ts on total lin ea r growth
and dry w eigh t accu m u lation .

T h e se sym ptom s appeared on plants of the

m en tion ed trea tm en ts in the follow in g order:

- N a fter 4 w eeks, 5 N and 5 P

a fter 6 w e ek s, - K and 5 K<S04> a fter 9 w eek s, 5 K(C1) and - F e after 10
w e ek s, -M g , - Mn, - B and - Cu a fter 1 1 w eek s, - P, - Ca, - Zn after 12
w eek s, and - Mo a fter 14 w e ek s.
It w as im p o ssib le to grow p lan ts without any nitrogen for longer than
th ree w e ek s.

M agnesium d eficien cy sym ptom s appeared m ore quickly on

plan ts w ith high p o ta ssiu m treatm en ts than on th ose treated with m agnesium
d eficien t solu tion .
V is ib le d e fic ie n c y sym ptom s on plants, as d escrib ed , w ere a sso c ia te d
w ith the follow in g nutrient le v e ls in the p e tio les:

nitrogen . 51%, phosphorus

. 10%, p o ta ssiu m .47%, calcium .26%, m agnesium . 14%, m anganese 18 ppm,
boron 14 ppm, copper 28 ppm, and zin c 11 ppm.
M any sig n ifica n t in tera ctio n s betw een nutrient e lem en ts w ere found.
In g en era l, m ajor e le m e n ts influenced the concentration of m inor elem en ts
in p e tio le s and ste m s m o re than m inor elem en ts influenced m ajor elem en ts.
High le v e ls of m ajor e le m e n ts in solution induced d eficien cy sym ptom s of
other e le m e n ts.
Som e o f the m o re str ik in g in tera ctio n s betw een elem en ts w ere as
follow s: (1) Low n itrogen in the solution resu lted in low phosphorus, low
boron, and high calciu m in p e tio le s . (2) High nitrogen produced high p h o s­
phorus and high boron p e tio le con ten ts.
cid ed with high m an gan ese and high iron.

(3) Low phosphorus in p e tio le s c o in ­
(4) V ery low le v e ls of copper and

125.

z in c w e r e found under d eficien t p o ta ssiu m conditions.

(5) A s p e c ific amount

of p o ta ssiu m w as needed to get adequate calcium and m anganese uptake.
(6) M ore p o ta ssiu m w as taken up when a s so c ia te d w ith the sulfate anion
than in com bin ation w ith ch lo rid e.
found in p e tio le s .

In the la tter c a se , m o re calcium w as

(7) High nitrogen, phosphorus, p otassiu m , m anganese

and boron in p e tio le s app eared under calcium d eficien t conditions.

(8) A

sig n ifica n t p o s itiv e c o r r e la tio n betw een total iron and m anganese in the
p e tio le s w as e sta b lish e d .

(9) Low phosphorus, p otassiu m , calcium , and

high m agn esiu m w e re a s so c ia te d w ith high m an ganese v a lu es in p e tio le s.
(10) M ore boron w as o b se rv e d in p e tio le s of plan ts trea ted w ith high le v e ls
of nitrogen, phosphorus, p o ta ssiu m , and low le v e ls of calcium than in those
plan ts tr e a te d w ith high le v e ls of boron in the solution.
With e x tr em e high m an ganese con cen tration s, m anganese toxicity
sym p tom s app eared on le a v e s .

The tra n sitio n point from non -toxic le v e ls

to to x ic le v e ls se e m e d to be w ith 2900 ppm m anganese p re se n t in the
petioles^.

126 .

LITERATURE CITED
A rch ibald, J. A. 1956. Grape nutrition stu d ies in Ontario. Paper p resen ted
at 36th annual m eetin g of A gr. Inst, of Canada, T oronto.
A skew , H. O. 1944. A c a s e of com bined p otassiu m and boron d e fic ie n c ie s
in g r a p e s. N. Z . Jour. S ci. T e c h ., Sec. A, 26: 146-152.
Babo A. von, and Ei M ach. 1910. Handbuch des W einbaues und der K ellerw ir tsch a ft. 1. Band, Weinbau. Paul P arey V erlag, Berlin.
Beattie, Jam es M. 1955. D e fic ie n c ie s of p otassiu m and m anganese red uce
grape y ie ld s . Ohio F arm and Home R es. N ov-D ee.
_________________ , and C. G. F o rsh ey . 1954. A su rv ey of the nutrition
e lem en t statu s of Concord g ra p es in Ohio. Proc. A m er. Soc. Hort.
S ci. 64: 2 1 -2 8 .
Bergman, E. L . , and A. L. K enworthy. 1956. Molybdenum d eficien cy and
n itra te to x ic ity on C oncord g ra p es. M ich. A gr. Expt. Sta. Quart.
Bui. 38 (4): 52 4 -5 2 7 .
. 1957. Seasonal variation of Concord
grape (V itis la b r u sc a ) p e tio le com p osition . Paper No. 308, p resen ted
at 54th annual m e e tin g of A m er. Soc. Hort. S c i . , Stanford, C alif.
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A gr. S ci. Bui. 353.
B ioletti, F . T. , and L. Bonnet.
R e s. 8: 3 8 1 -3 9 7 .

1917.

L ittle -le a f of the vine.

Jour. Agr.

Bonner, J . , .and A. W. G alston. 1952. P rin cip les of Plant Physiology.
W. H. F r e em a n and Company, San F r a n c isc o . 499 pp.
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P roc. A m er. Soc. H ort. S ci. 46: 246 -2 4 8 .
Branas, J . , and G. Bernon. 1954.
et V it. 142: 2 3 9 -2 4 3 .

Sur la ca ra n ce de bore.

Progr. Agr.

127.

Branas, J. , and G. Bernon. 1956. La caran ce de bore dans le s vignob les
fr a n c a is. P rogr. A gr. et V it. 43: 259 -2 7 0 .
Burghardt, H. 1956. B eitrage zum E isen-M an gan-A n tagonism u s der Pflanze.
F lo ra 143:(1) 1 -30.
s'

C ais, D. 1954. Note p r e lim in a ri su una m alattia della v ite risco n tra ta nel
V eron ese. R iv. V itic. E nol. VII/5: 164-167.
C handler, W. H. , D. R. Hoagland, and P. L. Hibbard. 1933. L ittle -le a f
or r o se tte of fruit tr e e s . Proc. A m er. Soc. Hort. S ci. 30: 70-88.
_____ _ ______________________________________________ . 1934. L ittle -le a f
or r o s e tte of fruit t r e e s . Proc. A m er. Soc. Hort. Sci. 3 2 :1 1 -1 9 .
Chapman, I. M. 1931. The rela tio n of iron and m anganese to c h lo r o sis
in p la n ts. New P h ytologist 30: 266-283.
C lore, W. J. 1951. L ittle -le a f or zin c d eficien cy on Concord grap es. Wash.
A gr. Expt. Sta. C ir. 136.
Coom bs, B. G. 1949. Z in c treatm ent of sultana v in e s.
S. A ust. 53: 5 9 -61.

Jour. Dept. Agr.

, and H. R. M. A llan. 1955. V ineyard nutrition on deep
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C alif.
Cooper, J. R. , and J. E. V a ile. 1939. R esp on se of A m erican grapes to
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Bui. 378.
Crawford, R. F. 1935. T h e cau se and control of c h lo r o sis in New M exico.
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DeKock, P. C. 1956. H e a v y m
N. S. X X /7 7 . January.

e ta l

to x icity and iron c h lo r o sis.

Ann. B o t.,

128.

D ille y , D avid R. 1957a. Growth and le a f com p osition in relation to the supply
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T h e s is for d eg ree of M .S ., M ichigan State U n iversity.
_______________ _•

1957b.

Unpublished data.

D ufrenoy, J. 1935. E ffets du zin c sur la c r o issa n c e des V itis.
Soc. Biol. 118: 156-158.

Compt. Rend.

Eaton, F . M. 1944. D eficien cy , to x icity and accum ulation of boron in plants.
Jour. A gr. R e s. 69: 237-2 7 7 .
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__________ . 1954. Starke Borm angelschUden an M osel und Nahe.
Weinbau, 19: Oct. 5.

W einberg

D eutscher

. 1955. U ntersuchungen liber den Kalium und M agnesiu m -gehalt
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G erb er, H. , and A. B ussm ann.
verfhrbungen bei R eben.

1955. U eber die v o rzeitig en H erbstSchw eiz. Z . O bst- u. Weinb. 64: 340-343.

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Kenworthy, A. L.

1957.

1950. The w ater culture m ethod for
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Unpublished data.

Kobayashi, A ., T. H osoi, and R. Isoda. 1955. Growth and yield of grapes
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______________ .

1957.

U npublished data.

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1957.

P erson al com m unication.

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Proc.

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1944.

Boron nutrition o'f g ra p es.

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Phytopath.

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A P P E N D I X

APPENDIX TABLE 1

NUTRIENT SOLUTION COMPOSITION, MAJOR ELEMENTS, USED IN EXPERI­
MENT I

T reatm ent

Ppm per

1 Liter*

N

P

K

Ca

Mg

so4

co3

Check

210

31

235

200

49

192

-

-N
5N

>5*>!»

1050

31
31

235
235

200
200

49
49

480
192

360

- P
P

210
210

155

235
235

200
200

49
49

192
192

60
60

- K
5 K (S 04)

5 K(Cl)

210
210
210

31
31
31

200
200
200

49
49
49

192
1342
192

- Ca
5 Ca (S 0 4)
5 Ca (Cl)

210
210
210

31
31
31

235
235
235

1002
1002

49
49
49

192
960
192

- Mg
5 Mg

210
210

31
31

235
235

200
200

5

-

_

1171
1171

-

-

243

-

Cl
-

-

-

-

-

-

-

849
-

-

300 532
1418

192
576

-

60

-

284

*M inor elem en ts as recom m ended by Hoagland and Arnon (1950).
* * O riginally no nitrogen, but 42 ppm added after 4 weeks in order to save
the plants.

135.

APPENDIX TABLE 2
NUTRIENT SOLUTION COMPOSITION, MINOR ELEMENTS, USED IN
EXPERIMENT I
Ppm p er 1 Liter*
T reatm en t
- Fe
5 Fe

Fe
-

25

Mn
.5
.5

B

Cu

Zn

Mo

.5
.5

.0 2
.0 2

.0 5
.0 5

. 01
.0 1

- Mn
5 Mn

5
5

2. 5

.5
.5

.0 2
. 02

.0 5
.0 5

.0 1
.0 1

- B
5 B

5
5

.5
.5

2. 5

. 02
.0 2

. 05
.0 5

. 01
.0 1

- Cu
5 Cu

5
5

.5
.5

.5
.5

. 10

. 05
. 05

. 01
. 01

- Zn
5 Zn

5
5

.5
.5

.5
.5

.0 2
. 02

.2 5

. 01
. 01

- Mo
5 Mo

5
5

.5
.5

.5
.5

.0 2
. 02

.0 5
. 05

-

-

*M ajor e le m e n ts u sed in the sam e amount a s for check;
s e e Appendix T able 1.

-

. 05

co

136.

o
c

CM

CM

CM CM

CMCM

O *5
feCOio
E ’

2
w

S
w
H
w

CM CM CM

CM CM CM

co

UO

CM CM

CO

co

Is

X

o

O
CO

E ^

m

uo

cm

2

CM

CM
u S

Dh

o

bo ^

§

cT'

[x ,

%

H

oc/3

m

uo

m

m

m

m

m

in

in

UO

uo uo uo

CM

22

H
£
W

fe,

HH

o

o

CM

p

CM

cd

c tf

O

H

D
£

°*3
$ 3
u •

Ix
O

W

W
H

2

eS

X
w

Ok
5j

CM
H
2

03

§ fe
O
W
w
&
<4 W

G ~
cd 2 ;
u ^

Is

in
cm

o

co

CM

"co

uo

uo

m

m

UO uo

cd

i". uo uo
'"f

uo
. in

CM

r-

uo uo

U

2
0

H

P

co

8X

2

L°

W

X
X

D

O 2

fe
HH
a
w

01
c/3
2:

oI—I
H
D
J
O

C/3
X
u

oH

C/3

ofe
C/3

fe

CM

CM

cm

.

C/3 LO
CM .
X

22

^C M

E
X
E
O
X

CM

uo

UO

uo uo

uo

uo

uo uo uo

uo uo

*
uo

CM

u 2
co
§ 2

uo

in

uo

UO uo

uo

uo

UO

03

H

CO
o
d
u

d
H
H

CD
E
■(->
cd

CD
X

E-h

M
O
a)

JS

U

3O ^CT?
C/3 U

r*
E
r-“*
<!

z:

2
i

uo

EX CX
1 UO

XXX
1

uo uo

o &

cn U

w ' ^
cd cd cd
U U U
O uo
i U

bO b0

£ s
I

uo

30. 9 grams CaSO^ added directly to nutrient solution.

fe

co

Minor elements stock solution see Appendix Table 4.

CO

o
o

137.

APPENDIX TABLE 4

STOCK SOLUTION COMPOSITION, MINOR ELEMENTS, USED IN PREPARING
NUTRIENT SOLUTIONS FOR EXPERIMENT I
Grams per
T reatm ent

MnCl2 ’ H2 O

1

Liter*
ZnSC>4

H3 BO4

CUSO4 - 5H20

. 900

1.430

.0 4 0

.108

.02

.0 4 0
.0 4 0

.108
.108

.02

4. 500

1 .430
1.430

*- B
5 B

.9 0 0
. 900

7.150

.0 4 0
.0 4 0

.108
.108

- Cu
5 Cu

.9 0 0
.9 0 0

1.430
1.430

.200

.108
. 108

- Zn
5 Zn

.9 0 0
.9 0 0

1.430
1.430

.040
.0 4 0

.5 4 0

.02

- Mo
5 Mo

.9 0 0
. 900

1.430
1.430

.0 4 0
.0 4 0

.108
.108

.10

All m a jo rs)
- Fe
)
5 Fe
)
- Mn
5 Mn

. * F errou s lactate, Fe(C 3 H5 0 3)2- 3H2 0 ,
as a separate stock solution.

12

H2 MoC>4 * 2 H2 O

.02

.02
.02

.02

‘.02
.0 2

--

. 956 grains per liter made up

138.

APPENDIX TABLE 5
INFLUENCE OF TREATM ENTS OF EXPERIMENT I ON DRY WEIGHT ACCUM­
ULATION AND TOTAL LENGTH GROWTH
T reatm en t
R oots

D ry W eight (G ra m s)
Stem s
P e tio le s L eaf Blades

Total

L inear
Growth (Cm)

C heck

2 3 .6 5

25. 60

4. 30

37. 15

9 0 .7 0

625

- N
5 N

16. 45
11. 85

7. 15
13. 95

1. 50
2 .6 5

13. 20
2 7 .4 0

38. 30
55. 85

109
336

- P
5 P

26. 55
1 2 .4 5

12. 90
18. 90

2 .6 0
3 .2 0

23. 00
2 7 .7 5

65. 05
61. 40

212
623

- K
5 K (S 0 4)
5 K (Cl)

1 1 .4 5
19. 55
9 .8 5

7 .0 0
19. 20
8 .8 5

1. 10
3 .7 0
2. 00

15. 90
31. 05
1 8 .5 0

35. 45
73. 50
3 9 .2 0

186
490
229

- Ca
5 Ca (SO ^
5 Ca (Cl)

2 1 .6 5
15. 80
19. 40

1 2 .4 0
1 0 .4 5
1 1 .2 0

2 .2 5
1 .7 5
1 .7 5

25. 20
18. 50
2 1 .0 0

61. 50
46. 50
53. 35

223
315
229

- Mg
5 Mg

1 5 .8 0
21. 35

22. 15
7. 15

3. 30
1 .2 5

36. 50
16. 40

7 7 .7 5
46. 15

518
178

- Fe
5 Fe

35. 05
24. 45

28. 75
17. 70

5. 11
3 .4 0

3 7 .6 5
29. 55

106. 55
75. 10

458
354

- Mn
5 Mn

27. 65
32. 35

19. 50
20. 80

3 .8 0
3. 60

29. 15
ai.25

80. 10
8 8 .0 0

464
468

- B
5 B

24. 70
2 8 .2 5

17. 75
2 4 .1 0

3. 15
4. 15

23. 00
3 0 .4 5

6 8 .6 0
86. 95

362
516

- Cu .
5 Cu

3 9 .6 0
34. 90

20. 05
24. 30

3. 05
4. 90

2 9 .2 5
36. 95

91. 95
101 .0 5

372
506

- Zn
5 Zn

2 9 .2 5
36. 05

18. 10
24. 85

2 .8 0
3. 70

2 8 .4 5
35. 05

78. 60
99. 65

390
482

- Mo
5 Mo

3 3 .2 0
49. 35

18. 85
26. 65

3. 35
4. 25

2 9 .4 5
37. 50

84. 85
1 1 7 .7 5

420
472

LSD 5%
LSD 1%

14. 11
19. 17

14. 07
19. 12

2 .5 7
3. 48

1 7 .2 4
2 3 .4 1

2 1 .0 7
2 8 .6 4

252
343

139.

APPENDIX TABLE 6
INFLUENCE OF TREATM ENTS OF EXPERIMENT I ON DRY WEIGHT ACCUM'
ULATION CONVERTED INTO PER CENT OF TOTAL DRY WEIGHT
Per Cent of Total Dry Weight

T reatm en t

Shoot/R oot
Ratio

R oots

Stem s

C heck

2 6 .2 9

28. 05

4 .7 1

40. 95

2 .8

- N
5 N

4 3 .8 6
21. 32

1 8 .2 9
24. 96

3 .8 0
4. 74

34. 05
49. 98

1. 3
3. 7

- P
5P

43. 77
20. 38

1 7 .9 8
29. 33

3 .8 4
5. 20

3 4 .4 1
4 5 .0 9

1. 3
3 .9

- K
5 K (S 0 4)
5 K (Cl)

3 2 .2 5
28. 10
24. 38

1 9 .7 5
25. 00
23. 07

3. 11
4. 68
5. 16

44. 89
42. 22
47. 39

2. 1
2. 6
3. 1

- Ca
5 Ca (S 0 4)
5 Ca (Cl)

3 5 .2 9
32. 54
36. 78

19. 97
23. 03
20. 31

3 .5 7
4. 10
3. 13

41. 17
4 0 .3 3
39. 78

1 .8
2. 1
1. 7

- Mg
5 Mg

20. 35
46. 35

2 8 .4 5
1 5 .4 3

4. 24
2. 70

46. 96
35. 52

3 .9
1 .2

- Fe
5 Fe

3 1 .5 9
3 2 .7 6

2 7 .4 3
2 3 .4 8

4. 96
4. 56

36. 02
3 9 .2 0

2 .2
2. 1

- Mn
5 Mn

33. 91
36. 62

2 2 .0 3
2 3 .6 4

4. 30
4. 09

3 0 .2 6
3 5 .6 5

1 .7
1. 7

- B
5 B

3 5 .4 7
34. 60

2 5 .8 4
26. 10

4. 60
4. 54

3 4 .0 9
3 4 .7 6

1.8
1. 9

- Cu
5 Cu

43. 08
36. 42

2 1 .8 2
23. 32

3 .3 1
4. 54

3 1 .7 9
35. 72

1. 3
1. 7

- Zn
5 Zn

3 7 .2 0
3 7 .8 6

2 2 .8 9
23. 94

3 .5 9
3. 57

36. 32
3 4 .6 3

1 .7
1 .6

- Mo
5 Mo

41. 04
4 4 .8 0

21. 04
21. 96

3. 75
3. 28

34. 17
30. 96

1 .4
1. 3

LSD 5%
LSD 1%

1 1 .8 0
16. 03

6 .8 0
9 .2 4

1. 53
2. 08

8. 55
1 1 .6 2

P e tio les

L eaf Blades

'

140.
APPENDIX TABLE 7
INFLUENCE OF TREATM ENTS OF EXPERIMENT I ON PETIOLE COMPOSITION
T reatm en t
N

P er Cent of Dry W eight
p
K
Ca
Mg

Fe

Ppm of Dry Weight
Mn
Cu
B

Zn

Check

1 .0 6

. 37

5 .7 2

. 99

.4 5

35

63

56

24

46

- N
5N

. 51
5. 19

. 21
.6 6

2 .2 4
1. 57

1.4 1
.4 6

.3 1
. 37

40
55

116
159

47
195

18
26

37
44

- P
5P

. 62
1. 41

. 10
1. 02

4. 65
4. 18

1. 48
. 46

.3 0
. 37

45
35

247
83

16
242

18
28

46
49

- K
5 K (S 0 4)
5 K (Cl)

1. 14
1. 18

. 75
. 68
. 52

. 12
9 .8 6
8. 88

.8 6
.6 4
.7 6

.9 9
. 14
. 14

70
35
45

268
44
52

45
229
198

11
23
20

25
42
56

2. 37
- Ca
5 Ca (S 0 4) 1 .0 9
5 Ca (Cl)
1. 18

1. 04
. 37
.4 2

6. 80
4. 51
2. 82

.2 6
. 96
2 .5 5

.5 2
.4 2
. 37

50
45
55

201
76
1^1

196
67
37

22
21
15

36
42
62

- Mg
5 Mg

1. 31
_*

.4 2
. 70

4. 92
4. 62

1 .6 7
. 58

.0 6
. 74

60
40

85
174

48
152

30
22

56
32

- Fe
5 Fe

. 95
1. 01

.2 9
.2 9

5. 13
4. 71

1. 48
1. 62

.6 5
.4 9

35
40

75
89

42
55

29
23

59
49

- Mn
5 Mn

1. 12
1 .0 4

. 32
. 37

5. 77
5. 59

1. 24
1. 21

. 50
.5 0

35
90

18
323

26
59

23
23

46
53

- B
5 B

1 .2 5
1. 08

. 42
. 35

5. 50
5. 32

.8 5
1 .5 4

. 46
.4 6

45
50

102
77

14
174

26
23

46
57

- Cu
5 Cu

1 .0 3
1. 12

. 33
.2 9

5. 35
5. 33

1. 43
1. 53

.4 8
. 54

50
35

63
42

24
38

29
30

42
49

- Zn
5 Zn

1.01
1. 05

.4 3
.4 5

5. 24
6. 33

1. 10
. 96

.5 5
. 46

45
50

82
46

33
118

24
28

48
42

- Mo
5 Mo

1 .0 6
1.01

. 28
.3 3

4. 85
5. 15

1 .7 9
1. 78

.5 3
. 54

35
50

84
94

21
58

22
22

64
47

* >:<

. 14
. 19

.4 2
.5 7

. 13
. 18

N. S.
N. S.

83
113

48
65

N. S. N. S.
N. S. N. S.

LSD v5%
LSD 1%

* N itrogen a n a ly s is o m itted due to shortage of plant m a teria l.
** A n a ly sis of v a ria n ce om itted due to m any m is s in g v a lu es.

141.
APPENDIX TABLE 8
INFLUENCE OF TREATM ENTS OF EXPERIMENT I ON STEM COMPOSITION

T reatm en t

N

Per Cent of Dry W eight
P
Ca
K
Mg

Check

1. 05

. 26

2. 37

- N
5 N

.6 3
2. 10

. 16
.3 0

1. 00
1 .8 7

- P
5P

.6 2
. 99

. 11
.4 0

- K
5 K <S04)
5 K (Cl)

1 .2 3
. 91
1. 07

- Ca
5 Ca (S 0 4)
5 Ca (Cl)

.6 2

Fe,

Ppm of D ry Weight
Mn
Cu
B

Zn

.2 7

40

36

45

22

34

.8 0
.3 7

.21
. 19

50
70

79
104

19
53

20
29

19
32

. 97
2. 52

. 77
. 36

.2 3
. 19

60
50

137
41

11
64

22
36

29
25

.2 6
. 26
.2 6

. 27
2. 61
2 .6 8

.4 3
.4 4
.4 9

. 33
. 18
. 19

100
40
40

122
28
31

26
41
29

28
22
22

33
28
31

1 .5 0
1. 10
.9 5

.2 5
.2 4
.2 4

1 .6 7
1. 86
1 .0 0

.2 5
. 91
1 .5 6

. 38
.2 8
.2 2

80
45
45

154
63
68

45
32
25

32
19
21

33
23
22

- Mg
5 Mg

. 91
1. 11

.2 5
. 32

1. 53
1. 70

. 97
.4 5

.0 7
.4 5

55
45

83
95

17
47

22
24

50
31

- Fe
5 Fe

.8 3
.9 6

. 19
.2 3

1. 65
2 .0 4

.7 2
. 79

.2 7
.2 6

40
45

44
40

18
26

24
24

39
35

- Mn
5 Mn

.9 1
1 .0 4

.2 1
.2 2

1 .9 8
1. 99

.7 2
.6 1

.2 7
.2 5

35
50

23
94

36
20

20
24

28
44

- B
5B

1. 12
1 .0 0

.2 3
.2 2

1. 56
1 .8 6

. 52
.6 3

.2 4
.2 4

45
50

59
47

11
55

23
30

20
41

- Cu
5 Cu

.8 6
.8 9

.2 1
.2 4

1 .4 2
1. 97

.7 9
.6 9

.2 6
.3 2

45
45

44
34

50
24

20
22

38
31

- Zn
5 Zn

. 92
. 93

.2 2
.2 3

1. 57
1. 69

.6 6
. 53

.2 8
.2 6

35
60

49
39

21
21

24
24

45
37

- Mo
5 Mo

.8 9
. 90

.2 1
.2 3

1. 69
1. 75

. 67
.8 5

.2 5
.3 9

45
55

50
56

26
28

31
38

36
48

LSD 5%
LSD 1%

.2 4
. 33

.1 1
. 15

. 58
.7 9

.2 8
. 38

.0 6
.0 9

N. S.
N. S.

34
47

31
35

N. S.
N. S.

17
24

APPENDIX TABLE 9
NUTRIENT SOLUTION COMPOSITION OF EXPERIMENT II
Ca+Mg
K

K

1

1 ,6

2

Ppm p er 1 Liter*
Ca
Mg

so4

Cl

117

100

24

96

--

3 .2

117

301

73

288

--

3

5 .3

23

100

24

96

4

6 .3

23

100

49

192

5

7 .4

23

100

73

288

--

6

8 .4

23

100

97

384

--

7

9 .6

23

200

24

96

--

8

1 0 .6

23

200

49

192

--

9

1 1 .7

23

200

73

288

--

10

12 .7

23

200

97

384

--

11

13.8

23

301

24

120

--

12

1 4 .9

23

301

49

216

--

13

1 5 .9

23

301

73

312

14

1 7 .0

23

301

97

408

--

15

18.1

23

401

24

120

177

16

1 9 .2

23

401

49

216

177

17

2 0 .2

23

401

73

312

177

18

2 1 .2

23

401

97

408

177

T reatm ent

*Nitrogen, phosphorus, iron, boron, zinc and molybdenum was the sam e
a s for the check treatm ent of Appendix Table 1 with the exception for
m anganese 1 ppm and copper . 03 ppm.

143.

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145.

APPENDIX TABLE 12
INFLUENCE OF TREATMENTS OF EXPERIMENT II ON DRY WEIGHT ACCU­
MULATION CONVERTED INTO PER CENT OF TOTAL DRY WEIGHT

T reatm en t

R oots

Per Cent of T otal D ry Weight
Stem s and
P e tio le s
L eaf
B erries
Blades

Shoot/R oot
Ratio

1
2

50. 45
41. 15

21. 17
25. 44

2. 14
2. 44

26. 24
30. 97

.9 8
• 1 .4

3
4
5
6

25. 72
26. 53
3 1 .9 7
34. 24

26. 97
3 1 .7 9
23. 56
2 5 .8 5

3. 00
2 .8 5
2. 99
2 .5 0

44.31
3 8 .8 3
4 1 .4 8
3 7.41

2 .9
2 .8
2 .1
1 .9

7
8
9
10

2 7 .6 5
24. 12
2 2 .6 7
3 0 .5 5

27. 26
34. 12
3 3 .6 7
29. 50

3. 39
2 .7 7
2 .8 8
2 .7 7

4 2 .7 0
3 8 .9 9
4 0 .7 8
3 7 .1 8

2 .7
3 .2
3 .4
2 .3

11
12
13
14

21. 97
25. 34
2 2 .8 1
29. 18

33. 07
33. 22
3 1 .0 2
29. 64

3. 12
2 .9 2
3. 28
3. 03

4 1 .8 4
3 8 .5 2
4 2 .8 9
38. 15

3. 6
3 .0
3 .4
2 .4

15
16
17
18

2 7 .1 0
2 7 .0 0
26. 42
2 8 .7 4

29. 90
31. 34
3 3 .2 0
2 8 .1 1

2 .9 8
2. 87
2. 93
2 .9 5

40. 02
3 8 .7 9
3 7 .4 5
40. 20

2 .7
2 .7
2 .8
2 .5

LSD 5%

N .S .

N. S.

N. S.

N. S.

146.

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148.

APPENDIX TABLE 15
SIGNIFICANCE OF F VALUES IN 4x4 FACTORIAL PORTION OF EXPERIMENT II
F - V alu es for
E lem en t

Plant P arts

N itrogen

P e tio le s
S tem s

Phosphorus

P otassium

C alcium

P e tio le s
S tem s

Calcium

__

M agnesium

Interaction
Calcium /M agne sium

_

_

NS

NS
NS
NS

>’fi
*

P e tio le s
S tem s

NS

P e tio le s
Stem s

sjs>;<
❖

NS
NS

NS
NS

##

NS
NS

NS
* >]«

NS
NS

P e tio le s
Stem s

NS

NS

P e tio le s
Stem s

NS
**

NS
NS

##

M anganese

P e tio le s
Stem s

NS

NS

NS
NS

Boron

P e tio le s
Stem s

*

NS
#

Copper

P e tio le s
S tem s

NS

NS

P e tio le s
Stem s

NS
NS

NS
NS

M agnesium

Iron

Z inc

--

Not d eterm in ed

* S ign ificant 5% le v e l
** S ign ificant 1% le v e l

NS

NS
NS

149.

APPENDIX TABLE 16
INFLUENCE OF MANGANESE LEVELS OF EXPERIMENT III ON DRY WEIGHT AC­
CUMULATION AND TOTAL LENGTH GROWTH
D ry W eight (Gram s)
T reatm en t

R oots

Stem s

B erries

P e tio les

L eaf
Blades

Total

L inear
Growth
(Cm)

Check
(1. 0 ppm Mn)

6 0 .2 3

1 2 .8 5

1 2 .4 3

2. 55

3 1 .3 3

119. 39

192

12. 5 ppm Mn

5 6 .2 5

29. 90

8 .4 6

5. 65

50. 10

150. 36

658

25 ppm Mn

63. 10

3 8 .8 0

12. 86

7 .2 5

65. 60

187.61

610

NS

NS

NS

2. 90

15. 35

40. 36

LSD 5%

1 8 2.83

150.

APPENDIX TABLE 17
INFLUENCE OF THE MANGANESE LEVELS OF EXPERIMENT III ON DRY WEIGHT
ACCUMULATION CONVERTED INTO PER CENT OF TOTAL DRY WEIGHT

T reatm en t
R oots

Per Cent of T otal Dry Weight
S tem s and B erries
L eaf Blades
P etio les

Shoot/R oot
Ratio

Check
(1. 0 ppm Mn)

5 0 .4 5

21. 17

2 .1 4

26. 24

12. 5 ppm Mn

37 .4 1

25. 51

3. 76

33. 32

1 .7

25 ppm Mn

3 3 .6 3

2 7 .5 4

3 .8 6

34. 97

2 .0

NS

NS

NS

NS

LSD 5%

.9 8

APPENDIX TABLE 18
INFLUENCE OF MANGANESE LEVELS OF EXPERIMENT III ON PETIOLE
COMPOSITION

T reatm en t

N

Per Cent of D ry Weight
P
Ca
K
Mg

Ppm of Dry Weight
Fe
Mn
Cu
B

Zn

Check
(1. 0 ppm Mn)

2. 18 .3 1

5 .8 6

2 .4 0

.6 5

52

371

22

22

50

12. 5 ppm Mn

1. 33 .2 9

3 .4 1

1. 72

.41

65

1840*

22

21

36

25 ppm Mn

1 .4 5

.2 6

3 .1 4

1 .7 2

.4 3

97

3067*

13

21

34

NS

NS

NS

NS

NS

28

965

NS

NS

NS

LSD 5%

*A ccu ra cy of sp ectro g ra p h ica l a n a ly sis for m anganese at th ese le v e ls m ay
be q u estion ab le.

152.

APPENDIX TABLE 19
INFLUENCE OF MANGANESE LEVELS OF EXPERIMENT III ON STEM
COMPOSITION
m

llC dllllC U L

Per Cent of D ry W eight
_
N
P
K
Mg
Ca

Ppm of Dry Weight
Fe
Mn
Cu
B

Zn

Check
(1. 0 ppm Mn) 1 .2 4

.1 9

.7 4

.5 7

. 18

34

91

9

21

15

12. 5 ppm Mn

1 .1 7

.2 5

1. 45

.8 4

. 17

46

265

15

17

17

25 ppm Mn

1 .1 3

.2 7

1 .2 8

.9 8

.2 0

62

658

12

17

21

LSD 5%

NS

NS

NS

NS

NS

NS

NS

NS

NS

NS