GROWTH AND LEAF CCMPOSIUON lN
RELATEQN TO THE SUPPLY GF CHLORIDE AND
SULFATE ANIGNS TO CHERRY, APPLE,
PEACH AW GRAPE FLAME

Thesis {or fits Degree of M. 5.
MICE'ESAN S'MYE UNWERSITY
David Ross Diiley
195‘?

 

THESIS

GROI~.TH AND LEAF COII‘TPOSITE ON IN RELATION TO TIE
SUPPLY OF CIHJOEIDE AID SULFATE ANIOI-JS TO

CHERRY, APPLE, PEACH AND GRAPE PLANTS.

BY

DAVID ROSS Dlm

AN ABSTRACT

Submitted to the College of Agriculture of Michigan
State University of Agriculture and Applied Scimce
in partial fulfillment of the requirements
for the degree of

EASTER OF SCIENCE
Department of Horticulture

1957

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A}X)I‘Ovedooo.000.000.00.0000000000000000.o 100.00....

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David Ross Dilley Abstract
ABSTRACT

The specific influence of chloride and sulfate anions, used singly
and in combination, upon growth and leaf composition of Kontmorency cherry,
Delicious apple, and Elberta peach, and Concord grape was studied by the
means of nutrient solution methods.

Three levels of chloride and sulfate were employed in a factorial
design giving nine treatment combinations without altering the metallic
cation content of Hoagland's nutrient solution.

Growth was measured as linear growth of shoots and dry weight of
various plant parts - leaves, shoots, fibrous roots, and trunk. Increase
in total dry weight was calculated. Leaves from.each tree and stems from
each grape plant were analyzed for twelve elements: nitrogen, phosphorus,
potassium, calcium, magnesium, iron, boron, manganese, copper, zinc,
sulfur and chlorine.

Increasing the supply of chloride decreased all measurements of
growth for all creps and produced leaf symptoms on peach and grape.

Increasing the sulfate supply did not adversely affect the growth of
apple, peach, and grape; but depressed growth of cherry trees.

Increasing the supply of chloride and sulfate together depressed
growth in proportion to the chloride level.

The chloride content in cherry leaves and grape petioles was directly
pr0portional to supply. Apple and peach leaves contained more chloride

when supplied with a medium chloride level. Increasing the sulfate level

David Ross Dilley Abstract

apparently reduced chloride absorption for all crOps.

Sulfur content in all plants increased with supply in the absence of
chloride. High chloride levels depressed sulfate absorption for all crops.

fioderate and high chloride levels depressed potassium absorption in
cherry, apple and peach while high levels of chloride increased potassium
in grape. Sulfate did not reduce potassium absorption in any crop.

Increasing the level of chloride in the solution increased nitrogen
in peach and grape; manganese in cherry, apple and peach; c0pper and zinc
in grape; and decreased; potassium in cherry, apple and peach; calcium.in
cherry and peach; magnesium in peach and grape; and boron in cherry.

Increasing the sulfate level in the solution increased nitrogen in
peach; phosphorus in cherry and peach; iron in peach and grape; boron in
cherry, apple and peach. Higher levels of sulfate decreased calcium in
cherry, apple and peach; and zinc in peach and grape.

Chloride and sulfate in combination increased nitrogen in peach and
phoSphorus in cherry and decreased calcium.in cherry and peach more than
when applied separately. Iron content in apple and zinc content in grape
was depressed more when chloride and sulfate were supplied in combination
than when applied singly.

Although growth depressions for all crOps with increasing chloride
or sulfate in the nutrient solution was of fine same magnitude, this could
not be attributed solely'to the concentration of chloride in the leaves.
The low or high level of certain elements induced by chloride or sulfate
levels were not considered inadequate nor toxic for growth according to

field observations. However, the relative pr0portions of certain elements

ii

David Ross Dilley Abstract

in the leaves induced by unequal anion supply and absorption may have been
a factor causing reduction in growth. Conversely, the accumulation of
certain ions in the leaves may have been the result of reduced growth

and not necessarily the result of absorption as influenced by chlorides.

GROWTH AND LEAF COMPOSITION IN RELATION TO
THE SUPPLY OF CHLORIDE AND SULFATE ANIONS

TO CHERRY, APPLE, PEACH AND GRAPE PLANTS

By

DAVID ROSS DILLEY

A THESIS

Submitted to the College of Agriculture of Michigan
State University of Agriculture and Applied Science
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE
Department of Horticulture

1957

ACKNOWLEDGEMENTS

The author wishes to express his sincere appreciation to Dr. A. L.
Kenworthy, for assistance in planning the experiment and in the preparation
of the manuscript; to Drs. R. L. Carolus, G. P. Steinbauer and K. Lawton
for editing the manuscript and to Dr. E. J. Benne and his staff for carrying
out the chemical analyses.

The author is grateful to the International Mineral and Chemical Company
whose financial assistance made this study possible.

Special acknowledgement goes to the author's wife, Marion, and to Dr.

F. G. Teubner for their constant encouragement and assistance.

TABLE OF CONTENTS

Page
INTRODUCTION ............................................ 1
REVIEW OF LITERATURE .................................. 2
MATERIALS AND METHODS ................................. 11
RESULTS .................................................. 14
Growth
General Observations ............................... 14
Increase in Total Dry Weight ........................ 20
Dry Weight of Fibrous Roots ......................... 22
Linear Growth of Shoots ............................. 23
Leaf Composition
Nitrogen. .......................................... 25
Phosphorus ......................................... 26
Potassium ........................................ ‘. . 27
Calcium ........................................... 28
Magnesium ........................................ 29
Iron ............................................... 30
Boron .............................................. 31
Manganese ......................................... 32
Copper ............................................ 33
Zinc ............................................... 34
Sulfur .............................................. 35
Chlorine ............................................ 36
DISCUSSION ................................................ 37
SUMMARY. . ............................................... 45
LITERATURE CITED ........................................ 48

APPENDIX .................................................. 53

INTRODUCTION

Depletion of potassium reserves in the soil through cropping eventually
leads to the addition of potassium fertilizers to maintain profitable crop
yields. In certain instances the use of quantities in excess of a thousand
pounds per acre are common. When applications reaching this magnitude
are used consideration of the associated "inert ingredients" or anion supplied
with potassium becomes increasingly important

Recognizing the harm ful effects of "inert ingredients" or anion many to-
bacco and blueberry producers no longer include potassium chloride in their
fertilizer programs due to adverse side-effects of the associated chloride
ion. Other crops such as beets can tolerate large amounts of chloride with-
out hurting crop yields.

Potassium fertilizers are becoming more generally used to satisfy the
requirements of fruit crops in most production areas. The use of greater
quantities of potassium fertilizers in fruit plantings may eventually expose
detrimental effects of the "carrier” components. Some components may be
beneficial, some detrimental, while still others may not influence plant
growth and quality.

The purpose of this experiment was to determine the specific effects of
chloride and sulfate anions, two common "carrier" components in potassium
fertilizers, as related to vegetative growth and leaf tissue composition of

Montmorency cherry, Delicious apple, and Elberta peach trees, and Con-

cord grape plants.

REVIEW OF LITERATURE

The importance of sulfur as a plant nutrient is generally known and
accepted. Chlorine, on the other hand, has been considered nonessential.
However, beneficial effects of chloride have been reported by Eaton (13) in
the nutrition of cotton and tomatoes. Young (60), Lipman (34), and Schmal-
fuss (51) report beneficial effects of chloride with a wide number of plants.
Recently, Broyer (6) established the essentiality of chloride in the nutrition
of higher plants. Arnon (2) reports intact green cells can carry on photo-
synthesis without the participation of chloride. However, once the cells are
broken, there is a rapid light induced deterioration of some cellular sub-
stance essential for photosynthesis. Chloride ion protects this substance
against inactivation while the intact cell accomplishes this in some other
manner.

While chloride has been found beneficial in various concentrations to
certain crops, other crops have been injured by chloride. Kelly (29) and
Reifenberg (47) found citrus trees were injured by a chloride concentration
above 200 p. p. m. in irrigation water. Kelley and Thomas (28) observed
excess chloride in the soil caused scorching and defoliation of citrus leaves.
Tip-burn and mottling of citrus leaves occurred in a nutrient solution con-
taining 710 p. p. m. of chloride, according to Chapman and Liebeig (9). Haas
(20) reported that healthy orange leaves contained 0. 48 per cent chloride
(dry weight basis) whereas leaves from injured trees contained 3. 48 per

cent chloride.

Boresch (5) found the use of chloride fertilizer blighted the edges of
currant leaves. Similar reports relate chloride accumulation to leaf scorch
in avocado, Haas (18) and pecan, Harper (22).

Deterioration of grape vines growing on saline soil was reported by
Ravikovitch and Bidner (46) due to high chloride content found in leaves from
affected vines (2. 47 per cent on dry weight basis) as compared to low chloride
concentrations (0.15) per cent) found in the leaves from healthy vines. Thomas
(55) reported that grapes growing on saline soil were afflicted with premature
opening of buds, small leaves and fruit, defoliation, and, in extreme cases,
faulty fruit set. Marginal and interveinal necrosis were symptoms of chlor-
ide toxicity appearing on leaves.

Jacobs et. a1. (25) found the addition of 150 pounds per acre of potassium
chloride injurious to leaves on young orange shoots, whereas the same rate
of potassium as sulfate was not injurious.

Parups (42), working with Montmorency cherry, observed a reduction
in tree growth resulting from an excess of either sulfate or chloride in the
nutrient solution. However, chloride reduced growth far more than sulfate
at the same concentration (670 p.p. m. ). High levels of chloride also re-
sulted in leaf scorch while similar levels of sulfate did not result in leaf
injury.

Hayward and Long (23) found tip-burn and mottling of peach leaves re-
sulted from growing trees in a high chloride nutrient solution. Both chloride
and sulfate reduced tree growth.

Magistad et. a1. (36) reported chloride to be more toxic than sulfate, in

isomotic concentration, with several vegetable crops.

Strong (54) studied the effect of calcium chloride used for dust control
along Michigan highways. He observed slight to extreme leaf scorch on
several shade trees during the summer months where run-off water con-
taining calcium chloride penetrated the soil in which the trees were growing.
Conifers were found more susceptible to injury than deciduous trees. Rudolfs
(49) found maple, birch, chestnut, and oak (in order of increasing tolerance)
susceptible to chloride injury.

The process of ion accumulation by plants has received considerable
attention during the past twenty years. Overstreet and Dean (40) have
summarized the accepted concents thusly:

"The process of ion absorption by plants requires expenditure
of energy. No ion accumulation occurs in the absence of respira-
tion and other metabolic activities. When metabolic activity is in-
hibited, accumulation of anions and cations is likewise inhibited.
The absorption process is an exchange process. Cations are ab-
sorbed in exchange for hydrogen ions and anions are absorbed in
exchange for hydroxyl or bicarbonate ions which are released to
the culture medium. The evidence indicates that no ion passes in
or out of a healthy plant except by exchange for another ion. Ion
accumulation is to a large extent selective. Due to the exchange
character of the process, anions can enter the plant independently
of cations and vise versa. All ions are not absorbed at the same
rates. In general potassium, ammonium, rubidium, and cesium
(monovalent cations) are rapidly accumulated while calcium, mag-
nesium and barium (divalent cations) are more slowly accumulated.
The anions nitrate, bromide. and chloride are usually rapidly ab-
sorbed. The anions sulfate and phosphate more slowly absorbed;
and anion bicarbonate is apparently not taken up at all. In concen-
trations below 0. 005 N, the rate of accumulation of an ion is de-
pendent on its concentration of the culture medium. "

It is obvious that the sum of anions plus cations must be balanced in the

plant to maintain electrical neutrality. Lundegardh (35) proposed that the

U1

uptake of anions creates the absorption potentially for the uptake of cations
in attaining this neutrality. Bear and Prince (4) established the concept of
cation constantancy in alfalfa and concluded that if one ion was absorbed to
a lesser extent, some other ion or ions would be absorbed to a greater ex-
tent to keep the sum of ions constant. The dissapation of formation of or-
ganic acids has been reported by Ulrich (S) to be the balancing mechanism
in maintaining cation-anion equivalency.

The literature pertaimng to the inter—relationship of anions and cations
as related to plant growth and composition is voluminous. Specific rela-
tionships as found for a particular crop may not hold true for the same crop.
and least of all for different crops, under a wide range of environmental con—
ditions. Many instances where chloride and sulfate anions have been found
to influence plant composition have been reported.

According to Leonard et. al. (33) sulfate depressed the absorption of
nitrate in sweet potatoes.

Carolus (8) working with beans found that sulfate, phosphate, nitrate,
and chloride did not detrimentally influence the uptake and utilization of cal-
cium. Buchner (7), on the other hand, found chloride enhanced absorption
of calcium in buckwheat and beans, especially when the ammonium form of
nitrogen was present.

Parups (42) studied anion balance in Montmorency cherry. He observed
that as chloride and sulfate concentrations were increased in the nutrient

solution the potassium content of leaves was decreased. However, chloride

exerted a far greater depressing effect than sulfate. Chloride was found to
enhance calcium and magnesium absorption whereas sulfate depressed the
absorption of these ions. Chloride and sulfate accumulation in leaves in-
creased with increasing concentration of these ions in the nutrient solution.

Cooper and Gortner (11), in their work with grapefruit, found leaf potas-
sium and calcium was reduced by high chloride while magnesium was not
affected. 1

Harper (22), found potassium accumulation in pecan leaves was not re-
duced by the presence of a high chloride content in the soil.

Haas and Reed (21), working with lemons, studied the absorption of chlor-
ide from various salts and reported that a greater quantity of chloride was
accumulated in leaves from potassium and calcium salts than from sodium or
magnesium salts.

Gilbert et. a1.(17) found that tung leaves absorbed more potassium when
supplied with monovalent anions than when supplied with divalent or polyvalent
anion sources. The anions studied were chloride, nitrate, sulfate, tartrate
and pectate.

Overstreet et. al. (41), working with excised barley roots, found calcium
absorption directly proportional to calcium chloride concentration up to
O. 200 N. Potassium absorption, however, increased with increasing potas-
sium chloride up to O. 08 N after which absorption dropped off markedly.

In a long term study of potassium chloride versus potassium sulfate in

grape nutrition, Vinet (58) concluded, after the third year of high applica—

tions, potassium sulfate proved to be a better source of potassium than
potassium chloride. The chloride associated with the high rates of appli-
cation, decreased the efficiency of potassium utilization while sulfate did
not.

Kenworthy (31) found grapes and peaches absorbed more potassium from
soils supplied with potassium sulfate than from soils supplied with potassium
chloride. The responses appeared to be related to soil type.

On the other hand, Skinner (53) found potassium chloride as good as
potassium sulfate as a fertilizer for cotton.

Buchner (7) found high chloride content reduced the ratio of inorganic
cations to inorganic anions in buckwheat and beans. Pierce and Appleman
(43) reported a similar response with several vegetable crops. In contrast
to those reports, Kretschmer et. al. (32) found that increasing the chloride
or sulfate concentration of the nutrient solution did not alter the ratio of
cations to anions with several of the same vegetable crops.

Cooper and Gortner (11) related the manifestation of chloride toxicity
to an excess accumulation of chloride over a total cation increase.

Rathje (45) working with lower green plants, observed reduced potas-
sium absorption as chloride, and to a lesser extent sulfate, accumulated
and attributed this to the acidifying effect of chloride on plant cells.

The dissipation or formation of malic acid, in the presence of excessive
anions or cations respectively, was thought to be a major factor in total

anion-cation balance in plants according to Ulrich (57).

Jacobson (26) and Jacobson and Ordin (27) reported that unequal absorp-
tion of anions and cations by barley roots was reflected by changes in or-
ganic acid content of roots. Greatest changes were observed in malic acid
content (the predominant organic acid present). Excessive cation accumula-
tion derived from a O. 005 N potassium bicarbonate solution, increased malic
acid content. Excessive anion accumulation, derived from a 0. 005 N calcium
bromide solution, decreased malic acid content.

Anderson et. al. (I) advised against the use of potassium chloride as a
fertilizer for tobacco. Chloride, being very mobile in the soil and easily
accumulated by tobacco, exerted a deleterious effect on burning quality of
tobacco. Sulfate, nitrate and carbonate of potash, being less easily absorbed
by tobacco, exerted a far less adverse effect on burning quality and were
therefore deemed better sources of potassium. Burning quality of tobacco
has been related to high potassium and high organic acid (especially bicar-
bonate, malic and citric acid) content of the leaf.

Garner et. al. (15) found a high chloride concentration associated with
dissipation of malic acid content in tobacco leaves. Since organic acids are
the major component of the buffer mechanism of plant cells conditions affect—
ing their dissipation could have direct bearing on pH control within the cell
and the associated protoplasmic activity.

Dunn and Nylund (12) studied the influence of fertilizers on specific
gravity of potatoes. In general, high specific gravity was associated with

high starch content and high cooking quality in potatoes. They concluded.

that potash fertilizers containing chlorides caused a marked reduction in
specific gravity, and the reduction was related directly to application. Sul-
fate of potash did not reduce specific gravity of the potatoes, indicating that
the chloride anion was responsible for increasing the water content of potatoes.

Baslavskaya (3) reported that potato leaves from plants receiving chloride
contained more starch than leaves from plants not receiving chloride.

Gauch and Eaton (16) found barley plants had higher starch and sugar con-
tent when grown in saline sand culture. This suggested that the accumulated
salts interfered with cellular activities associated with carbohydrate utilization.

Riesenauer and Colwell (48) found chloride absorption by tobacco plants
directly related to the hydrogen ion concentration of the culture medium.
Calcium, nitrogen and sulfate reduced chloride absorption.

Nielson and Overstreet (38), working with barley roots, reported a de-
crease in potassium absorption was directly related to the hydrogen ion con-
centration of the culture medium. Calcium decreased the inhibiting effect
of hydrogen ion on potassium uptake.

Overstreet, Jacobson and Handley (41) reported that calcium stimulated
potassium absorption in barley roots in high potassium solutions but inhibited
potassium absorption in low potassium solutions.

As early as 1902 Wheeler and Hartwell (61) recognized the toxic effect of
chloride to certain plants and reported that liming the soil could counteract
this toxicity. Piland and Willis (44) reported similar beneficial results of

liming to counteract chloride toxicity in certain plants.

Buchner (7) working with buckwheat and beans found chloride absorption
was favored in the presence of the ammonium form of nitrogen as compared

to the nitrate form.

10

11

MATERIALS AND METHODS

Specific effects of chloride concentrations, sulfate concentration and
chloride-sulfate combinations on the growth and leaf composition of cherry,
apple, peach and grape were studied by means of nutrient solution methods.
To single out specific effects of chloride, sulfate and chloride-sulfate com-
binations three concentrations of chloride (0, 355, 655 p. p. m.) and sulfate
(192, 384, 624 p. p. m.) were employed in a 3 x 3 factorial design Cochran
and Cox (10).

To obtain the nine nutrient solutions containing the concentrations of
chloride and sulfate desired, Hoagland's (24) nutrient solution was used
and the metallic cation content kept constant.1 This was accomplished by
using different amounts of various C. P. grade chemicals dissolved in dis-
tilled water and varying the nitrogen source between ammonium and nitrate
nitrogen (Appendix Tables 1 and 2). Conductivity readings were taken on each
solution with a Solu-bridge (model DR-15) and the pH of the solutions was
measured with a Beckman pH meter (model H2).

The following kinds of fruit plants were used: one-year-old Montmorency

cherry trees (Prunus cerasus budded on Prunus Mahaleb; one-year—old De-

 

 

licious apple trees (Pyrus malus budded on Pyrus malus); one-year-old

 

 

Elberta peach trees (Prunus persica budded on Prunus persica. var. Lovell);

 

 

 

To maintain Hoagland's metallic cation content it was necessary to reduce
the concentration of chloride and sulfate to 497 and 574 p. p. m. respectively
in the treatment containing the high levels of each.

12

and one-year-old own-rooted Concord grape (Vitus labrusca) cuttings.

 

Twenty seven plants of each kind of fruit were selected. The trees were
pruned uniformly and divided in three size groups according to weight after
pruning. Trees with approximately the same weight were put in the same
replicate, thus making three replicates for each treatment. ‘ The grape plants
were selected to provide the same weight after pruning to two buds. Green
and dry weights of representative plant parts were obtained from a separate
lot of plants and the per cent dry matter calculated. This value was used to
obtain initial dry weight of the experimental plants.

The trees and rooted cuttings were planted in 12-inch clay pots in the
greenhouse on January 28, 1956. The pots were previously painted with an
asphaltum compound to prohibit root contact with the surface of the clay
pots. Coarse quartz sand was used as the growing media.

The pots were numbered and arranged in three randomized blocks for each
crop on four greenhouse benches. Application of nutrient solutions commenced
one week after planting. Each plant received one quart of solution every third
day during the first month, and one quart every second day during the second
month. Later as growth demanded, daily application of solution was necessary.
The plants received bi-weekly and later tri-weekly flushing with distilled water
to avoid excessive salt accumulation in the pots. At the beginning of the third
month, and extending for two weeks, nutrient solution was withheld from plants

receiving the high chloride treatment. Distilled water was substituted in an

effort to bring the plants back into growth. During this two week period of

13

distilled water application, growth was renewed and the regular schedule of
nutrient application was re-established.

The plants were harvested on May 21, 22, 23 and 25. Linear growth of
shoots, dry weight of leaves, shoots, fibrous roots, large roots and trunk was
obtained and increase in total dry weight was calculated. Dry weight of fibrous
roots was obtained by passing dried, crumbled roots through two standard
seives (sizes U. S. 20-840 microns opening and U. S. 45-350 microns opening).
Those roots passing through the smaller seive were weighed as fibrous roots.

Leaves from each tree and stems1 from grape were analyzed in the Agri-
cultural Chemistry Department Laboratory. Nitrogen was determined by the
Kjeldahl method, potassium by flame photometer, phosphorus, calcium, mag-
nesium, iron, manganese, boron, copper and zinc were determined spectro-
graphically. Chloride was determined by a modified method of Samson (50).

Sulfate was determined gravemetrically in accordance with the A. O. A. C.

method (39).

 

1
Stem analysis rather than petiold analysis was necessary due to insuffic-

ient petioles from certain treatments. However, petioles were used for
chloride and sulfate determination.

14

RESULTS

Growth
Data pertaining to increase in total dry weight, dry weight of shoots, leaves
fibrous roots, and linear growth of shoots for the four crops can be found in
Appendix Tables 3, 4, 5 and 6.

General Observations

 

Within one month after the application of nutrient solution was initiated,
visual differences in linear growth of all crops due to differential sulfate and
chloride treatment were observed. Linear growth of all crops was depressed
as the chloride concentration in the nutrient solution increased. Of the four
crops, apple appeared most sensitive and peach least sensitive to the high
(665 p.p. m.) chloride concentration. No visual differences in linear growth
due to sulfate concentration were observed for any crop until the end of the
second month. However, at the end of the second month, linear growth of
cherry had been depressed slightly from both medium and high sulfate levels.
No response to sulfate concentration was observed for apple and grape. Peach
trees seemed to benefit from a medium (384 p.p. m.) sulfate supply. The
leaves took on a dark green cast and the shoots appeared more vigorous than
on trees receiving less sulfate (192 p.p. m. ). Visual linear growth differences
due to sulfate and chloride in combination were expressed mainly as a chlor-
ide effect for all crops. Differences in growth of all crops due to treatment

became more pronounced with time. Figures 1 and 2 show the response of

'l HESlS

 

apple and peach trees to the treatments. The response of cherry and grape

plants were similar to that shown for apple and peach.

Leaf symptoms developed on peach trees receiving high chloride solutions
by the end of the second month. The symptom appeared first on trees receiv-
ing high chloride (665 p.p. m.) and low sulfate (192 p. p. m. ). Similar, though
far less intense, symptoms appeared later on trees receiving high chloride
and medium sulfate (384 p.p. m. ). The terminal leaves were afflicted first.
As the new leaves unfolded a chlorosis developed beginning at the base and
extending up the margin of the leaves. This chlorosis soon gave way to a
marginal necrotic scorch as the leaves enlarged. Leaves so afflicted abscissed
readily (Figure 2 - upper right hand corner). The general leaf color was pale
green on trees receiving high chloride, compared to the dark green leaves on
trees receiving no chloride. Slender stems and small leaves were also a re-
sult of high chloride treatment.

Leaf symptoms also developed on peach trees receiving the medium chlor-
ide (315 p.p. m.) - high sulfate (624 p.p. m.) solution. The terminal leaves
were most severely damaged. Interveinal chlorosis started at the base and
extended to the tip of terminal leaves. The interveinal chlorosis gave way to
a necrotic "shot-hole" effect accompanied by considerable crinkling near the
midrib. Similar, though less intense, symptoms resulted from the medium
chloride - medium sulfate solution.

Grapes receiving medium chloride - high sulfate solution developed leaf

symptoms during the fourth month as shown in Figure 3. Leaves along the

THESIS

 

THESIS

 

 

 

Figure 1.
Influence of chloride and sulfate supply on the growth of Delicious apple trees.
From left to right --- increasing chloride supply from 0—315—665 p. p. m.

From top to bottom —-- increasing sulfate supply from 192-384-624 p. p. m.

16

THESIS

 

 

THESIS

 

 

Figure 2.
Influence of chloride and sulfate supply on growth of Elberta peach trees.
From left to right - -- increasing chloride supply from 0-315-665 p.p. m.

From top to bottom - - - increasing sulfate supply from 192-384-624 p. p. m.

17

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THESIS

 

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Figure 3.

Influence of medium chloride (355 p. p. m.) and high sulfate (624 p. p. m.) on
Concord grape.

18

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19

middle and extending upward including those on the terminal portion of the
shoot showed interveinal chlorosis. The chlorosis soon developed into ne-
crotic blotches covering a considerable area of the leaf. The medium chlor-
ide - medium sulfate treatment gave similar, though less intense, symptoms.
Leaf symptoms did not develop on cherry or apple from chloride or sul-
fate treatment. However, the leaves of both crops were much smaller on

trees supplied with chloride than on trees not receiving chloride.

THESIS

 

THESIS

HONTMORENCY CHERRY

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Increase in total dry weight. (see figure opposite)

 

Increasing the chloride concentration from O to 665 p. p. m. with 192 p.p. m.
of sulfate reduced dry matter accumulation in cherry trees. The 384 and 624
p. p. m. sulfate levels accentuated the depressing effect of increased chloride
concentration. Increasing the sulfate supply from 192 to 384 p. p. m. to trees not
receiving chloride reduced dry matter accumulation but further increase in sul-
fate supply did not.

As the chloride concentration was increased from 0 to 315 p. p. m. with 192
p. p. m. sulfate dry matter accumulation in apple trees was reduced slightly where-
as increasing chloride to 665 p. p. m. drastically reduced it. The 384 and 624
p. p. m. sulfate concentrations accentuated the depressing effect of increasing
chloride supply. Increasing the sulfate supply to trees not receiving chloride
did not influence dry matter accumulation but when chloride was included it was
depressed.

Increasing the chloride supply to peach trees receiving 192 p. p. m. sulfate re-
duced dry matter accumulation with each increment in chloride. The 384 and 624
p.p. m. sulfate levels accentuated the depressing effect of chloride. Supplying
384 p. p. m. of sulfate without chloride enhanced dry matter accumulation in peach
trees while further increase in sulfate supply to 624 p. p. m. reduced it.

Grape plants tolerated 315 p. p. m. of chloride when supplied with 192 or 384
p.p. m. of sulfate but the 665 p. p. m. chloride concentration drastically depressed
dry matter accumulation regardless of the sulfate supply, A stepwise reduction

in increase on total dry weight occurred as the chloride supply was increased with—

THESIS

 

 

 

 

in the 624 p.p. m. sulfate concentration. Increasing the sulfate supply from
192 to 624 p.p. m., regardless of chloride supply, did not effect dry matter

accumulation in grape plants.

21

THESIS

 

 

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22

Weight Of fibrous I'OOtS (see figure opposite)

Increasing the chloride concentration from 0 to 665 p. p. m. with 192 p. p. m.
sulfate reduced the dry weight of fibrous roots from cherry trees. The higher
sulfate levels accentuated the depressing effect of the 355 p.p. m. chloride con-
centration. Increasing the sulfate supply from 192 to 384 p.p. m. in the absence
of chloride reduced fibrous root growth while further increase in sulfate supply
did not induce additional growth depression.

As chloride was increased from 0 to 315 or 355 p. p. m. with 192 or 384 p. p. m.
of sulfate root growth of apple trees was not affected but when supplied with 624
p. p. m. of sulfate fibrous root growth was reduced. Increasing the chloride
supply to 665 p. p. m. reduced root growth within all sulfate concentrations. The
depressing effect of 355 and 665 p. p. m. chloride was accentuated as the sulfate
supply increased. Increasing the sulfate supply from 192 to 624 p. p. m. to trees
not receiving chloride did not affect fibrous root growth of apple trees.

The 315 or 355 p.p. m. chloride concentrations did not affect root growth of
peach trees receiving 192 or 384 p.p. m. of sulfate. The 665 p. p. m. chloride
level drastically reduced root growth with all sulfate concentrations. At the 624
p. p. m. sulfate level the depressing effect of 355 p.p. m. of chloride became more
pronounced.

Increasing the chloride concentration from O to 665 or 500 p.p. m. within‘ any
sulfate level reduced root growth in grape plants. Increasing the sulfate supply
enhanced root growth when supplied without chloride. At the 355 and 665 p. p. m.

chloride concentrations increasing the sulfate supply did not markedly affect

growth of grape roots.

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23

Linear growth of shoots (see figure opposite)

Increasing the chloride concentration from 0 to 665 p. p. m. with the 192 p. p. m.
sulfate level reduced shoot growth of cherry trees. Increasing the chloride level
within the 384 and 624 p. p. m. sulfate concentration tended to accentuate the de-
pressing effect of the 355 p. p. m. chloride concentration. Linear growth of

cherry shoots was reduced as much by the 384 p. p. m. sulfate as the 624 p. p. m.
sulfate level or the 315 p. p. m. chloride concentration.

A s the chloride supply to apple trees receiving 192 p. p. m. of sulfate was in-
creased from 0 to 665 p. p. m. a reduction in shoot growth resulted. Increasing
the Chloride supply depressed linear growth of apple shoots more as sulfate was
increa sed in the nutrient solution. Increasing the sulfate supply in the absence
of chloride or with 665 p. p. m. of chloride did not influence growth of shoots.
However, increasing the sulfate supply to trees receiving 355 p. p. m. of chloride
reduced growth.

The 315 p. p. m. chloride concentration did not influence linear growth of peach
8h00ts when supplied with 192 p. p. m. of sulfate while the 665 p. p. m. chloride con—
centration markedly reduced growth. As the chloride supply was increased with
the 384 or 624 p. p. m. sulfate levels the growth depression due to chloride be-
came more pronounced. Increasing the sulfate supply from 192 to 624 p. p. m. en-
hanced growth of peach shoots on trees not receiving chloride, depressed growth
when SuPplied with 355 p. p. m. of chloride, and did not significantly affect growth
When supplied with 665 p. p. m. of chloride.

Linear growth of grape shoots was not appreciably influenced by increasing

the Chloride supply from O to 315 or 355 p. p. m. within the 192 or 384 p. p. m.

HESI 5

an.

 

24

sulfate concentrations but drastically reduced growth when supplied with 624

p.p. m. of sulfate. The 665 p. p. m. chloride concentration did not greatly re-

duce growth unless used with 384 or 624 p. p. m. of sulfate. Sulfate when
supplied without chloride tended to increase growth but when chloride was

added sulfate accentuated the depressing effect of chloride.

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25

Leaf Composition
Data pertaining to the chemical composition of leaf tissue for the four fruit
crops can be found in Appendix Tables 7, 8, 9 and 10.

Nitrogen (see figure opposite)

Increasing the supply of chloride in the nutrient solution from O to 665 p.p. m.

had no effect on nitrOgen content in cherry leaves from trees supplied with 192 .1}
I
p.p. m. of sulfate. Though the differences were not significant increasing the
chloride concentration within the 384 p.p. m. sulfate and more so within the 624
p. p. m. sulfate levels tended to reduce nitrogen accumulation. Nitrogen content :1
if

in cherry leaves was not markedly influenced by increasing the sulfate supply
regardless of the chloride concentration.

. Apple leaves did not vary significantly in nitrogen content as the chloride
supply was increased from 0 to 665 p.p. m. regardless of the sulfate concentra-
tion. Leaves from trees supplied with increasing amounts of sulfate and no
chloride had nearly the same nitrogen content.

Increasing the chloride concentration from 0 to 665 p. p. m. within the 192
p. p. m. sulfate level increased the nitrogen content in peach leaves. As chloride
levels were increased with higher sulfate levels there was a greater increase in
nitrogen content than resulted from increasing the chloride supply within the 192

p.p. m. sulfate concentration. All sulfate levels in the absence of chloride gave

nearly the same nitrogen values.
Grape stems had a higher nitrogen content when the chloride concentration

”was increased within the 192 and 384 p.p. m. sulfate levels but not within the 624

p, p. m. sulfate concentration. Increasing the sulfate supply in the chloride had

no effect on nitrogen content in grape stems.

 

 

 

 

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26

Phosphorus (see figure Opposite)
Phosphorus content in cherry leaves increased then decreased slightly as the
chloride level in the nutrient solution was increased from 0 to 665 p. p. m. within

" the 192 p. p. m. sulfate concentration. As the chloride concentration was in-

creased from 0 to 665 p.p. m. within the 384 and 624 p. p. m. sulfate levels phos-
phorus content decreased. Cherry trees receiving increasing sulfate levels in

the absence of chloride increased with each increment increase in sulfate supply

but not when chloride was included in the nutrient solution.

Apple leaves showed no significant differences in phosphorus content but tended

to have more phosphorus in their leaves as the chloride supply was increased

from O to 665 p. p. m. within the 192 p. p. m. sulfate level. Increasing the chlor-

ide supply from O to 355 p.p. m. within the 384 and 624 p. p. m. sulfate levels

tended to increase phosphorus in apple leaves while the 665 or 500 p. p. m. chlor-

ide concentrations tended to reduce it. The sulfate supply did not significantly

affect the phosphorus content.

Peach leaves increased in phosphorus content as the supply of chloride in-

creased within all sulfate concentrations. Increasing the sulfate supply increased

phosphorus accumulation in peach leaves at the O and 355 p.p. m. chloride con-
centration but not at the 665 p.p. m. chloride concentration.
Grape stems tended to contain more phosphorus as the chloride supply was

increased from 0 to 665 p.p. m. within the 192 and 384 p.p. m. sulfate levels but

not at the 624 p. p. m. sulfate concentration. Sulfate supply did not markedly in-

II uence phosphorus content.

 

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27

Potassium (see figure opposite)
Cherry leaves decreased in potassium content as the chloride concentration

was increased from 0 to 665 p. p. m. within all sulfate levels. The 355 p. p. m.

chloride level exerted a greater depressing effect on potassium accumulation as

the sulfate supply was increased. Increasing the sulfate supply in the absence

of chloride or with 665 p. p. m. chloride did not markedly change the potassium

content while increasing the sulfate supply with 355 p. p. m. of chloride reduced

it.

Apple leaves contained less potassium as the chloride supply was increased
from 0 to 665 p. p. m. regardless of the sulfate concentration in the substrate.

Increasing the supply of sulfate exerted no influence on potassium accumulation

regardless of the chloride concentration.

Peach leaves decreased in potassium as the chloride supply was increased
from 0 to 665 p. p. m. within all sulfate levels. The depressing effect of 355

p. p. m. of chloride was accentuated as the sulfate supply was increased. In-

creasing the sulfate supply in the absence of, or with 665 p. p. m. of chloride did

not effect the potassium content in peach leaves.

Grape stems decreased slightly then increased in potassium content as the

chloride supply was increased from O to 665 p. p. m. within all sulfate levels.

At the higher sulfate levels these differences were less pronounced. In the ab-

sence of chloride, increasing the sulfate supply depressed potassium accumula-

tion. When chloride was added to the nutrient solution increasing the sulfate

supply accentuated the depressing effect of chloride on potassium accumulation.

1,3... ._

 

 

 

 

 

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28

Calcium (see figure opposite)

Cherry leaves decreased in calcium content as the chloride supply was in-
creased from 0 to 665 p. p. m. within all sulfate levels. Increasing the sulfate
supply in the absence of chloride decreased the calcium content but not as severely
as did the medium or high chloride levels when supplied with any sulfate concentration.

The calcium content in apple leaves tended to increase as the chloride supply
was increased from 0 to 665 p. p. m. within the 192 and 624 p.p. m. sulfate concen-
trations though the differences were not significant. Increasing the sulfate con-
centration from 192 to 624 p. p. m. in the absence of chloride reduced the calcium - 3.‘
content in apple leaves. The depressing effect of sulfate on calcium accumulation
was exerted regardless of the chloride concentration.

The calcium content in peach leaves was reduced as the chloride supply was
increased from 315 p.p. m. to 665 p. p. m. within the 192 p. p. m. sulfate concen-
tration. At the 384 or 624 p. p. m. sulfate levels increasing the Chloride supply
from O to 355 reduced calcium accumulation in peach leaves. As the sulfate
supply was increased from 192 to 624 p. p. m. concurrently with O or 355 p. p. m.
of chloride the calcium content was markedly reduced. The lowest calcium value
was observed at the 355 p. p. m. chloride - 624 p. p. m. sulfate concentrations.

Grape stems decreased then increased in calcium content as the chloride
supply was increased from 0 to 665 p. p. m. regardless of the sulfate supply. In-
creasing the sulfate supply with O or 255 p. p. m. of chloride reduced the calcium
content. The greatest reduction in calcium content in grape stems occurred

when 355 p. p. m. of chloride and 624 p. p. m. of sulfate were combined.

THES
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29

Magnesium (see figure opposite)

 

The magnesium content in cherry leaves decreased as the chloride supply was
increased from 0 to 315 or 355 p. p. m. within the 192 and 384 p. p. m. sulfate levels.
Increasing the chloride concentration from 315 to 665 p. p. m. within the 192 p. p. m.
sulfate level did not further reduce the magnesium content. With the 384 p.p. m.
sulfate concentration increasing the chloride supply from 355 to 665 p. p. m. in-
creased the magnesium content in cherry leaves while with the 624 p. p. m. sulfate
level it did not. As the sulfate supply was increased from 192 p.p. m. in the ab-
sence of chloride the magnesium content in cherry leaves was lowered.

Magnesium content in apple leaves increased as the chloride supply was in-
creased within the 192 and 384 p. p. m. sulfate concentrations. Within the 624 p. p. m.
sulfate level increasing the chloride supply increased the magnesium content at
the 355 p. p. m. chloride level. Increasing the sulfate supply from 192 to 624 p.p. m.
in the absence of chloride or with 665 p. p. m. of chloride did not markedly change
the magnesium content. Magnesium content was increased by supplying 624 p. p. m.
of sulfate with 355 p. p. m. of chloride.

Leaves from peach trees decreased markedly in magnesium content as the
chloride supply was increased from O to 665 p.p. m. regardless of the sulfate level.
Increasing the sulfate supply in the absence of chloride tended to decrease the mag-
nesium content but not when chloride was included in the substrate.

Increasing the chloride supply from 0 to 665 p.p. m. within the 192 and 624
P-P- m. sulfate levels decreased the magnesium content in grape stems. When
sulfate was supplied at the 384 p. p. m. level increasing the chloride supply did
n0t reduce magnesium. The quantity of sulfate supplied did not effect the mag—

nesium content regardless of the chloride concentration.

THE:

 

 

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30

_I_r_on_(see figure opposite)

Increasing the chloride supply from O to 665 p.p. m. within any sulfate level
did not significantly change the iron content in cherry leaves. Iron content in
cherry leaves increased as the sulfate supply was increased from 192 to 624
p.p. m. in the absence of chloride. When the sulhte supply was increased from
384 to 624 p. p. m. within the. 355 p. p. m. chloride concentration cherry leaves
accumulated more iron.

Apple leaves showed no great change in iron content as result of increasing
the chloride concentration from 0 to 665 p. p. m. within the 192 and 384 p. p. m.
sulfate levels. However, higher levels of chloride reduced iron when 624 p.p. m.
of sulfate was used. Increasing the sulfate supply from 192 to 384 p. p. m. did
not change the iron content in apple leaves while increasing the sulfate supply
from 384 to 624 p. p. m. in the absence of chloride markedly increased it. .

Iron content in peach leaves increased then decreased slightly as result of in-
creasing the chloride supply\ from O to 665 p. p. m. within the 192 and 384 p. p. m.
sulfate concentrations but in the 624 p. p. m. sulfate level it did not. In the ab-
sence of chloride or when supplied with 665 p. p. m. of chloride, increasing the
supply of sulfate from 192 to 624 p.p. m. increased iron content in peach leaves.

Increasing the chloride supply from 0 to 665 p.p. m. increased the iron con-
tent in grape stems more as the sulfate concentration was increased from 192 to
624 p.p. m. When sulfate was supplied without chloride no great change in iron
content in grape stems was observed. However, when chloride was included in
the nutrient solution increasing the sulfate supply from 192 to 624 p. p. m. in-

creased the iron content.

 

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31

_1§9_r_()__n (see figure opposite)

The boron content in cherry leaves did not change as the chloride supply was
increased from 0 to 315 p.p. m. w th the 192 p.p. m. sulfate concentration while
increasing the chloride supply to 665 p.p. m. markedly reduced it. With the 384
p. p. m. sulfate, increasing the chloride concentration reduced boron accumulation.

Increasing the sulfate supply from 192 to 624 p. p. m. in the absence of chloride in— '75-}
creased boron accumulation. The depressing effect of the 665 p. p. m. chloride con- '1‘}.
centration on boron accumulation was counterbalanced by increasing the sulfate supply.

Apple leaves tended to contain less boron as the chloride supply was increased
from 0 to 665 p.p. m. with the 192 and 384 p. p. m. sulfate concentrations. At
the 624 p. p. m. sulfate level boron increased markedly then decreased as the
chloride supply was increased from 0 to 500 p.p. m. Increasing the sulfate supply
from 192 to 624 p.p. m. in the absence of chloride, or when 665 p. p. m. of chlor-
ide was included, slightly increased the boron content while with 355 p. p. m. of
chloride it was increased markedly.

Peach leaves increased then decreased in boron as the chloride supply was in-
creased from 0 to 665 p. p. m. with the 192 "ml 624 p. p. m. sulfate concentrations.
Increasing the sulfate supply from 192 to 624 p.p. m. increased the boron content
in peach leaves regardless of the chloride level.

The boron content in grape stems increased as the chloride supply was increased
from O to 665 p. p. m. with the 192 p. p. m. sulfate level but had no effect with the
384 or 624 p. p. m. sulfate concentrations. Increasing the sulfate supply did not in-

fluence boron accumulation in grape stems regardless of the chloride concentration.

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32

Manganese (see figure opposite)

 

A linear increase in the manganese content in cherry leaves resulted when
the chloride supply was increased from 0 to 665 p. p. m. with the 192 and 384 p. p. m.
sulfate levels but not at the 624 p. p. m. sulfate level. Increasing the sulfate supply
from 192 to 624 p.p. m. reduced manganese accumulation only in the presence of
the 665 p. p. m. chloride concentration.

Apple leaves increased in manganese content as the chloride supply was in-
creased from 0 to 665 p. p. m. regardless of the sulfate supply. Increasing the
sulfate supply from 192 to 624 p. p. m. tended to reduce the stimulatory influence
of chloride on manganese absorption.

Peach leaves tended to increase in manganese as the chloride supply was in-
creased from 0 to 665 p. p. m. with the 384 and 624 p. p. m. sulfate concentrations.
Increasing the sulfate concentration increased manganese accumulation in peach
leaves only in the presence of the 665 p. p. m. chloride concentration. .

Manganese increased then decreased in grape stems as the chloride supply
was increased from O to 665 p.p. m. with all sulfate levels. Increasing the sul—
fate supply from 192 to 624 p.p. m. to grape plants not receiving chloride in—

creased the manganese content.. The stimulating effect of sulfate on manganese

accumulation was less pronounced as chloride was added to the nutrient solution.

 

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33

Copper (see figure opposite)
Increasing the chloride supply from O to 665 p.p. m. decreased then increased
the copper content in cherry leaves from trees receiving 192 p.p. m. of sulfate.
The exact reverse occurred when chloride was increased with the 384 and 624
p. p. m. sulfate levels. Increasing the sulfate supply from 192 to 624 p. p. m. in
the absence of chloride, or with 665 p.p. m. of‘chloride, depressed copper accum- WWI-w

ulation and increased it when 355 p.p. m. of chloride was included.

. 1'..." I.u.«_.. .

Copper content in apple leaves tended to be higher when chloride was supplied
at the 355 p. p. m. concentration. Increasing the sulfate supply tended to increase
the copper content though the differences were not significant.

Peach leaves did not vary significantly in copper‘content as the chloride
supply was increased from 0 to 665 p.p. m. regardless of the sulfate concentra-
tion. Increasing the sulfate supply from 192 to 624 p.p. m. in the absence of
chloride decreased then increased the copper content in peach leaves. When
supplied with chloride, however, increasing the sulfate supply did not increase
the copper content.

Increasing the chloride supply from 0 to 665 p.p. m. increased the copper
content in grape stems regardless of the sulfate concentration. However, when
supplied with 384 or 624 p. p. m. of sulfate, copper increased only when supplied
with 665 p. p. m. of chloride. In the absence of chloride increasing the sulfate
supply did not alter the copper content while increasing the sulfate within the

355 p. p. m. chloride concentration increased the copper content.

DELICIOUS APPLE

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34

Zing (see figure opposite)

Cherry leaves decreased in zinc as the chloride supply was increased from 0
to 665 p. p. m. regardless of the sulfate concentration. The 315 or 355 p. p. m.
chloride concentration did not markedly change zinc content while the 665 p. p. m.
concentration reduced it. Increasing the sulfate supply from 384 to 624 p. p. m.
reduced zinc absorption when supplied with 355 p. p. m. of chloride.

Zinc cmtent in apple leaves did not vary significantly as result of increasing '
the chloride supply from 0 to 665 p. p. m. regardless of the sulfate level. In-
creasing the sulfate supply from 192 to 624 p. p. m. in the absence of chloride or
when supplied with 355 p. p. m. of chloride tended to decrease zinc accumulation.

Increasing the chloride concentration from 0 to 665 p. p. m. within thel92
or 384 p. p. m. sulfate levels slightly increased then decreased zinc accumulation
in peach leaves. At the 624 p.p. m. sulfate concentration zinc accumulation de-
creased linearly as the chloride supply was increased. As the sulfate supply
was increased from 192 to 624 p. p. m. in the absence of chloride or when 665
p. p. m. of chloride was included the zinc content was not changed while including
355 p. p. m. of chloride decreased it.

Zinc content in grape stems was not markedly changed by increasing the
chloride supply from 0 to 355 p.p. m. regardless of the sulfate concentration. At
the 665 p.p. m. chloride concentration zinc accumulation was greatly increased.
Increasing the sulfate supply from 192 to 624 p. p. m. in the absence of chloride
01' when 355 p. p. m. of chloride was added did not change the zinc content while
increasing the sulfate with 665 p. p. m. of chloride counteracted the stimulating

effect of high chloride on zinc accumulation.

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35

_S_p_l_f_ur (see figure opposite)

Sulfur content increased then decreased in cherry leaves as the chloride con-
centration was increased from 0 to 665 p.p. m. regardless of sulfate supply. At
the high sulfate level, high chloride markedly depressed sulfur accumulation. In—
creasing the sulfate supply in the absence of chloride or when supplied with 355
p. p. m. chloride brought about a linear increase in the sulfur content in cherry
leaves. Chloride depressed sulfur accumulation when both were supplied in high
concentration.

Apple leaves increased then decreased in sulfur as the chloride supply was
increased from 0 to 665 p. p. m. especially when increased within the high sulfate
concentration. Increasing the sulfate supply from 192 to 624 p.p. m. increased the
sulfur content in apple leaves more when supplied with 355 p. p. m. of chloride
than when supplied without chloride or with 665 p. p. m. of chloride.

Increasing the chloride level from 0 to 665 p. p. m. did not change the sulfur
content in peach leaves from trees supplied with 192 p. p. m. sulfate. When supplied
with 384 p. p. m. of sulfate, chloride increased then decreased sulfur accumulation.
At the 624 p. p. m. sulfate level, increasing the chloride supply from O to 500 p. p. m.
decreased the sulfur content in peach leaves. Increasing the sulfate supply from
192 to 624 p.p. m. increased sulfur content in peach leaves more when supplied
with a moderate amount of chloride than with a high amount of chloride.

The high chloride level (665 p. p. m.) decreased and the moderate chloride
level (355 p. p. m.) increased sulfur content in grape petioles from plants supplied
with 384 and 624 p. p. m. sulfate while chloride did not influence sulfur content in

plants receiving 192 p. p. m. Of sulfate. Sulfur content in grape petioles increased

more with increasing sulfate supply in the presence of a moderate chloride con-

centration than in the absence of chloride or in the presence of a high level.

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36

Chlorine (see figure opposite)

Cherry leaves contained more chloride as the supply in the nutrient solution
was increased from O to 665 p. p. m. within the 192 and 384 p. p. m. sulfate levels.
In the presence of 624 p. p. m. of sulfate increasing the chloride supply increased
then decreased the quantity of chloride recovered in cherry leaves. Increasing
the sulfate supply to 624 p. p. m. decreased the chloride content in cherry leaves
from trees supplied with 51)") p. p. m. chloride.

The chloride content in apple leaves increased then decreased slightly as the
concentration in the nutrient solution was increased from 0 to 665 p. p. m. regard-

less of the sulfate level. The 384 and 624 p. p. m. sulfate concentrations reduced

the chloride content in apple leaves from trees supplied with moderate and high

chloride levels.
Peach leaves increased then decreased slightly in chloride content as the

Supply was increased from 0 to 665 p. p. m. The decrease in chloride content at

the high chloride level was accentuated as the sulfate supply was increased from
192 to 624 p. p. m. Increasing the sulfate supply from 192 to 624 p. p. m. reduced
the amount of chloride recovered in peach leaves from trees supplied with moder-
ate and to a greater extent high chloride concentrations.

The amount of chloride recovered in grape petioles was linear with supply in
the nutrient solution when supplied with 192 or 624 p. p. m. of sulfate. At the 384
p .p. m- sulfate concentration chloride content in grape petioles increased then de-
créaSed as the chloride supply was increased from 0 to 665 p. p. m. Increasing
the sldfate supply from 0 to 384 p. p. m. accentuated chloride accumulation in grape

t‘
pe loles from plants receiving the moderate chloride level. High chloride and high

Sulfat -
e In combination accentuated chloride accumulation.

THE

 

 

37

DISCUSSION

Although the concentration of essential elements, except chloride and sul-
fate, was maintained at a constant level, marked differences in growth and leaf
composition resulted. Since the supply of elements was kept constant, Variation
in growth and leaf composition can be contributed to specific anion effects.

Increasing the supply of chloride to a high level (665 p. p. m. ) resulted in an
On

apparent toxicity as measured by severe growth depression in all crops.

peach trees receiving a high chloride - low sulfate solution the leaves showed
marginal scorch -- -a symptom described by Hayward and Long (23) and others

(5), (18), (22), (42) as chloride toxicity.

Increasing the sulfate supply did not adversely effect growth of apple, peach,
and grape; but depressed growth of cherry trees.

Parups (42) reported that sulfate and chloride were absorbed by cherry
trees in direct proportion to their supply in the substrate. This was not found
to be true for all crops under the conditions of this experiment. Greater quanti-
ties of chloride were recovered in apple and peach leaves from the medium ra-
ther than the high cl-loride level. Sulfate appeared to reduce chloride absorption.
This effect was accentuated for all crops as the sulfate supply was increased in
the culture medium. The sulfur content in the leaves of all crOps increased with
increasing sulfate supply in the absence of chloride. As the chloride level in-
Crefilsed there was first a stimulation then a depression in sulfur accumulation
in the leaves of all crops.

It was quite apparent that the reduction in growth of all, and the development

 

 

 

 

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of visible toxicity symptoms in some, plants was a direct result of the chloride
concentration in the nutrient solution. Supplied with the same chloride concentra-
tion, the growth of all plants was depressed nearly the same magnitude. Since
chloride ions are freely absorbed, an accumulation Of chloride ion in the leaves
should have been related to growth. The highest chloride content found in cherry
leaves was 0. 4; in apple - 0. 24; in peach — l. 24; and in grape petioles - l. 82 per
cent of their dry weight. The chloride content in cherry and apple leaves was
considerably lower than found in peach leaves and grape petioles yet growth re-
sponse was very similar. The greatest growth depression occurred in cherry
trees when the chloride content in the nutrient solution was highest. This was
not true for apple, peach and grape.

Peach leaves having the highest and nearly the same chloride content, ex-
hibited two entirely different leaf symptoms, depending on the chloride - sulfate
supply in the substrate. Marginal scorch occurred with the high chloride - low
sulfate solution; but interveinal "shot-hole" necrosis occurred with medium chlor-
ide - high sulfate or medium chloride - medium sulfate solutions. The latter
two solutions developed blotchy-necrotic leaf symptoms on grape.

Inasmuch as the chloride content in the leaves of apple, peach, and petioles
of grape cannot be entirely responsible for growth depression, the effect of chlor-
ide may have been expressed as limiting the absorption or utilization Of some
other element(s) or necessary growth factor.

There was an inverse relationship between chloride and potassium in the

leaves. However. the chloride content in the leaves was not always highest

39

when potassium content was lowest in the leaves. High chloride content in the
nutrient solution always resulted in a lower potassium content of the leaves.
This would infer that the depressing effect of chloride On potassium accumula-
tion was exerted during absorption by roots. Overstreet et. al. (41) report de-
creased potassium absorption by barley roots as the concentration of potassium
chloride increased in the substrate. Potassium and chloride ions are absorbed
by roots by two distinctly different mechanisms and would therefore not be ex-
pected to compete with each other for metabolic "binding sites" according to
Epstein (14).

Several workers (11), (45), (58), have reported that the accumulation of
chloride cirectly or indirectly interfers with potassium accumulation in several
plants. In studies where the concentration of essential elements was kept con-
stant and the chloride level increased, Parups (42) found potassium content in
cherry leaves decreased markedly.

Boresch (5) theorized that while potassium may be present in adequate
amounts for normal plant requirements, there may not be a sufficient quantity
to combine with all the chlorine in the leaves. According to him, the excess
chloride is combined with calcium and an antagonism between potassium and cal-
cium produces the symptoms of potassium deficiency. This concept seems in-
adequate to explain the expression of symptoms on two counts; dissociation
phenemenon regarding calcium chloride salts could not effect tie up calcium be-
cause the chloride ion can be extracted quantitatively from leaf tissue by water;

and the chloride ion accumulates under conditions of excess supply in quantities

40

greater than. calcium absorption could account for.
Sideris and Young (52) conclude from their work with pineapple
"that the terminal leaf necrosis of pineapple is affected by changes

in the metabolic condition Of the tissues resulting from high chloride

concentration in association with potassium deficiency. Pineapple

plants amply supplied with potassium even in the presence Of high

chloride concentration failed to develop leaf necrosis. The chemical

differences in leaf tissue composition between plants with ample and

with deficient potassium are expressed mainly in the concentration

Of starch and sugars. It is possible that starch and sugars produced

with ample potassium increase the tolerance Of the cells to high chlor-

ide concentrations. "

The physiological response to chloride ion may be the result of enzyme
activation brought about by the increased acidity of the plant cells due to chlor—
ide. Large increases in acidity changes the acceleration to an inhibition of
enzyme activity according to Haas (19). The marked changes in the organic
acid metabolism in tobacco leaves, Anderson, et. al. (1) and barley roots
Jacobson and Ordin (27), induced by excessive anion accumulation may be re-
flected by acceleration or inhibition of essential enzyme systems related to
cellular activities associated with carbohydrate utilization.

To compare the leaf composition of crops grown in the greenhouse to leaf
composition of the same crops grown in the field is not completely valid. A
comparison of low, high and average values observed in the field to those ob-
tained in the greenhouse may, however, gove some insight to growth responses

that Occurred. Under field conditions Kenworthy (30) found the following average

composition in cherry, apple, and peach leaves.

41

 

Per cent of dry weight

 

N P K Ca Mg Mn Fe Cu B

 

Montmorency cherry 2. 83 . 267 l. 54 l. 91 . 74 . 0114 . 028 . 0055 . 0067

Jonathan apple 2.45 .232 1.65 1.58 .44 .0112 .021 .0010 .0057

 

Elberta peach 3.79 .243 1.56 1.92 .71 .0101 .023 .0016 .0060 m.“

When these values are compared to the values for the check treatment
(0-chloride and 192 p.p. m. sulfate) it is seen that the potassium values are 3 1. 1
considerably higher than those found in the field. With the exception of potassium, ~..:
the leaf composition of peach compares quite closely to field averages. The
composition of cherry leaves grown in the greenhouse varied considerably
above and below field averages - nitrogen was higher and phosphorus, boron,
iron and manganese were lower. Greenhouse apple leaves were higher in ni-

trogen and calcium but the remaining elements compared favorably with field

 

averages. A composition comparison of greenhouse grown grapes to those
grown in the field was not possible because stem analysis could not be compared
to petiole analysis available fOr field conditions.

Though chloride depressed potassium accumulation in cherry, apple and
peach the level was still considerably above averages considered deficient in
the field. The same was true for calcium in cherry, magnesium in peach and
cherry, and boron in cherry and apple. The low calcium content in apple and

peach leaves from high sulfate treatment was still above values considered de-

 

42

ficient in the field. The high sulfate - medium chloride solution reduced cal-
cium content in peach leaves to a low level which was still considered adequate
according to field data.

The tolerance of plants to chloride injury varies widely depending upon
species and prevailing climatic conditions. Environmental conditions which
favor high transpiration rates are conducive to salt injury on plants (such as,
squash, tomatoes and alfalfa) when supplied with high salt concentrations accord-
ing to Magistad et. al. (36). Neller (37) reported that plants growing in soils
high in organic matter can tolerate more chloride than plants growing in sandy
soils.

The chloride content in the soils varies widely depending upon the use of
fertilizers containing chloride, proximity to waters containing chloride, deposi—
tion of chloride salts during the soil forming process and the degree of leaching
that has progressed.

When fertilizer is applied to a soil and conditions conducive to leaching pre-
vail, loss of nutrients in drainage water is likely to occur. The proportion of
the component ions in the drainage water usually deviates from those added in
the fertilizer because of cation and anion exchange reactions in the soil. This
is well illustrated by the Observations of Volk and Bell (59) on lysimeters
filled with a loamy fine sand having a base exchange capacity of 3. 0 m. e. per
100 grams. Various sodium, magnesium, and potassium salts were applied to
different lysimeters. Though no calcium was applied it was the predominant
cation recovered in the drainage water. Only 1 per cent Of the potassium; 65

per cent Of the sodium; and 30 per cent of the magnesium applied was recovered

43

in the leacate. However, chlorides were recovered almost quantitatively;

about 50 per cent of the sulfates and almost twice as much nitrate was recovered
as was applied. There was practically no difference in the proportion or con-
tent of cations in the leachate, whether the salts added were chlorides or ni-
trates; but when sulfate salts were applied there was a marked decrease in the
loss Of calcium.

From these results of Volk and Bell (59) it would be infered that the chlor-
ide ion would not accumulate in soils. However, the ability of a soil to retain
chloride depends somewhat upon soil type and structure, Wilklander (62).
According to Toth (56) and Wilklander (62) acid clay soils retain more chloride
than the same soils with higher pH values.

The effect of chloride ion on promoting loss of nutrients in the soil, mainly
calcium, and its depressing effect on the absorption of certain elements, es-
pecially potassium, by fruit trees, may be of considerable importance. Fruit
trees growing on light soils deficient in these elements, under conditions of
limited rainfall, where loss Of chloride through leaching would be minimized,
quite possibly may be injured by large applications of fertilizers containing
chloride. The chloride ion, being freely mobile in soils with low exchange
capacities, may concentrate in the soil solution as the moist=~re supply is di-
minished by the trees. Fertilizer may be applied to a limited area around
fruit trees. If this rate of application to a few square feet is converted to an
acre basis, the application may amount to several thousand pounds per acre.
Relatively heavy applications of potassium chloride may approach the concen-

trations used in this experiment. However, the application is not renewed during

 

 

44

the season and any toxic effects of the relatively high chloride level would be
of short duration. This may account for the lack of visible symptoms of chlor-
ide toxicity under field conditions.

The buffering effect of soil organic matter, colloidal clay, and bicarbonate
on rendering plants more resistant to chloride injury coupled with loss of chlor-
ide through leaching reduces the hazard of using fertilizers containing chloride.
Beneficial effects of liming to reduce chloride injury according to Wheeler and
Hartwell (61) and Piland and Willis (44) further reduce the hazard of using chlor-
ide fertilizers. Avoiding the use of an ammonium form Of nitrogen when using
fertilizers containing chloride has been recommended by Buchner (7).

Potassium fixation does not appear to be related to the form in which it is
supplied. F ctors governing the adsorption Of chloride by soil colloids and
accumulation of chloride in plants, may result in less potassium ions accumula-
ted in plants supplied with the chloride form of potassium as compared to the
sulfate or carbonate forms. There are some indications that this occurs under
field conditions, Kenworthy (31), Vinet (58). However, the evidence does not
warrant avoiding the use Of potassium chloride as a fertilizer if used with

consideration to existing soil and climatic conditions and crop being grown.

45

SUMMARY

The specific influence of chloride and sulfate anions, used singly and in
combination, upon growth and leaf composition of Montmorency cherry, De~
licious apple, and Elberta peach, and Concord grape was studied by the means
of nutrient solution methods.

Three levels of chloride and sulfate were employed in a factorial design
giving nine treatment combinations without altering the metallic cation content
Of Hoagland's (24) nutrient solution. This was accomplished by using different
amounts of various chemicals containing the anions involved.

Growth was measured as linear growth of shoots and dry weight of various
plant parts - leaves, shoots, fibrous roots, and trunk. Increase in total dry
weight was calculated. Leaves from each tree and stems1 from each grape
plant were analyzed for twelve elements: nitrogen, phosphorus, potassium,
calcium, magnesium, iron, boron, manganese, copper, zinc, sulfur and chlorine.

Increasing the supply of chloride decreased all measurements of growth for
all crops and produced leaf symptoms of chloride toxicity on peach and grape.

Increasing the sulfate supply did not adversely affect the growth of apple.
peach, and grape; but depressed growth of cherry trees.

Increasing the supply of chloride and sulfate together depressed growth in

proportion to the chloride level.

The chloride content in cherry leaves and grape petioles was directly

 

1 See footnote page 13.

46

proportional to supply. Apple and peach leaves contained more chloride when
supplied with a medium chloride level. Increasing the sulfate level apparently
reduced chloride absorption for all crops.

Sulfur content in all plants increased with supply in the absence of chloride.
High chloride levels depressed sulfate absorption for all crops.

Moderate and high chloride levels depressed potassium absorption in cherry,
apple, and peach while high chloride levels favored potassium absorption in
grape plants. Sulfate did not reduce potassium absorption in any crop.

Increasing the level of chloride in the solution increased nitrogen in peach
and grape; manganese in cherry, apple and peach; copp er and zinc in grape:
and decreased; potassium in cherry, apple and peach; calcium in cherry and
peach; magnesium in peach and grape; and boron in cherry.

Increasing the sulfate level in the solution increased nitrogen in peach;
phosphorus in cherry and peach; iron in peach and grape; boron in cherry, apple
and peach. Higher levels of sulfate decreased calcium in cherry, apple and
peach; and zinc in peach and grape.

Chloride and sulfate in combination increased the concentration of nitrogen
in peach and phosphorus in cherry and decreased calcium in cherry and peach
more than when applied separately. Iron content in apple and zinc content in
grape was depressed more when chloride and sulfate were supplied in combination
than when applied singly.

Although growth depression for all crops with increasing chloride or sulfate

in the nutrient solution was of the same magnitude, this could not be attributed

 

Vina.

.f‘.

47

solely to the concentration Of chloride in the leaves. The low or high level

Of certain elements induced by chloride or sulfate levels were not considered
inadequate or toxic for growth according to field Observations. However, the
.relative proportions of certain elements in the leaves induced by unequal anion
supply and absorption may have been a factor causing reduction in growth.
Conversely, the accumulation of certain ions in the leaves may have been the F—T
result of reduced growth and not necessarily the result of absorption as in-

fluenced by chlorides.

 

10.

ll.

48

LITERATURE CITED

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49

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