THE INFLUENCE 0? SOIL CONDITIONS AND FERTILIZER
TREATMENTS OH THE GROV/TH, CHEMICAL COMPOSITIONt AITD
ENZYME ACTIVITIES Of SUGAR BEETS.

BY

James Tyson

A THESIS

Presented to the Committee on Advanced Degrees
of the Michigan State College of Agriculture and
Applied Science in Partial fulfillment of the Require­
ments for the Degree of Doctor of Philosophy.

Soils Department
East Lansing, Michigan
1929

ProQuest Number: 10008445

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ACKNOWLEDGMEN T

The writer wishes to express his appreciation
to the Chilean Nitrate of Soda Educational Bureau of
New York for their financial support a:id to Dr. M. M.
McCool, who suggested these investigations, A)r this
and many valuable suggestions concerning the details
of the exoenrnentation and for aid in preparing the
manuscript•

<t

71298

CONTENTS
Page
I*

Introduction

II.

Review of Literature

III. Methods of Analysis

IV.

1.

Minerals

2.

Sugar

3.

Enzymes

Experimental Results
1.

2.

3.

Seasonal Variations of Mineral Nutrients
a.

Rifle muck

b.

Hillsdale sandy loam

c♦

Miami loam

Variations of Mineral Nutrients due to fertilizers
a.

Hillsdale sandy loam

b•

Rifle muck

c.

Miami loam

d.

Berrien sandy loam

e.

Brookston silt loam

Influence of Soil Conditions and Fertilizer
Treatments on Sugai* Contant

4.

Influence of Soil Conditions and fertilizer
Treatments on Activities of Enzymes

V.

Discussion

VI•

Summary

VII. Literature Cited

INTRODUCTION
It is well known that certain soil conditions are
suitable for the production of certain crops, and that
other conditions may cause crop failures.

Even unpro­

ductive soils may be made highly productive by the addi­
tion of only one fertilizer constituent, whereas others
require more than one.

Conversely the addition of one or

two fertilizer salts may cause decrease in growth of
plants on even productive soils.
The soil conditions and fertilizer treatments which
cause such varying results in the growth of plants must
cause some physiological change in the intake of nutrients
them which influences their life processes and growth*
Evidently there are certain ratios of nutrient elements
which are optimum for the development of the plant and for
the functioning of the physiological processes within it.
The investigations reported In this paper were under­
taken to study the influence of soil conditions on the
growth of the beet root, on the ratios and percentages of
mineral nutrients absorbed by the leaves and roots of beets,
on the sugar contents of the beet roots, and on the enzyme
activities of the leaves of b e t s .

More and more importance

is being; attributed to enzymes by plant physiologists and
biochemists who believe "the mechanisms of those chemical
transformations taking place in the life processes of plants
and animals are affected principally by the actions of
enzymes."

REVIEW OF THE LITERATURE.
Literature dealing With the influences of nutrient
conditions on the growth and composition of plants, and
on the activities of enzymes

is so voluminous that it

is necessary to limit this review to those papers which
hear most directly upon the questions under considera­
tion.
Variations in the amount of mineral elements absorbed
by the plants have been reported by investigators since
after Humphrey Davy (l) drew attention to the fact that
mineral constituents were essential for the development
of plants.

Wolff (2) in a review of plant ashes states

that "the quantitative composition of one and the same
plant varies according to the soil upon which it is grown."
Plant physiologists attribute the intake of mineral
nutrients to the two processes, osmosis and imbibtion.
It is well known also that plant tissues exhibit selective
absorption towards minerals in the nutrient solution
surrounding them.

Loew (3) remarked " it is well known

that plants cultivated in the presence of more sodium
than potassium, nevertheless absorb a greater quantity of
the latter than of the former, and some plants grown on
soils rich in potassium will absorb almost no sodium".
The permeability of the cell walls of living organ­
isms, according to McDougal (4), is influenced by the action

-

8-

of salt ions in solution around them.

It is quite

probable that the different ions in solution change
the permeability of the cell walls in this way when
the soil solution is altered in any manner, and cause
changes in the rate and ratio of absorption of mineral
nutrients.

This undoubtedly is the reason why magnesium

is not readily absorbed by plants in the absence of
calcium, and is injurious, while in the presence of
calcium it is taken up and the life processes of the
plant proceed in a normal manner.
This toxic and antitoxic effect of salts was ob­
served by Loew (3) who stated that '’magnesium salts can
perform their nourishing functions only in the presence
of calcium, while in its absence they exert an injurious
effect.TT McCool (5) found that magnesium, sodium, and
potassium present in pure solutions were very toxic to
seedlings.

This toxicity was greatly reduced in full

nutrient solutions and in soil cultures.

Antagonism

resulted when certain cations were present together in
solution, among which were sodium and potassium, and
calcium and magnesium.

Calcium was the most effective

of all the substances studied.
Breazeale (6) reported the absorption of phosphoric
acid to be increased by the presence of nitrogen in the
nutrient solution, while the absorption of calcium was
not increased by the presence of nitrogen, phosphoric
acid, or potash.

Potassium absorption was increased by

the presence of other plant foods, especially nitrogen.

-

3-

Hoagland (7) grew bafley in water cultures and
found a marked absorption of nutrient elements at all
periods up to the final stages of growth, when suitable
concentrations of ions were maintained.

Intense absorp­

tion during the later stages of growth led to no import­
ant increases in crop yield, which seemed to be conditioned
in a large measure by a favorable supply and concentration
of nutrients during the early stages of growth.

The

percentage and total quantity of nitrogen and potassium
per plant was decidedly increased in tops with incieased
concentrations of the nutrient solutions,

Loew (3)

reported that plants absolutely require a certain minimum
of each nutrient mineral, and in most cases besides this
minimum they take up not only an excess of these various
compounds, but also substances which are perhaps useful,
but. not absolutely essential for plant functions.

This

surplus, he explains, depends upon the intensity of the
current of transpiration to a large extent.
Burd (8) described a progressively increasing
absorption of nitrogen, potassium, calcium, and magnesium
by barley, grown in soils under controlled moisture condi­
tions, ending about the time heads began to form, after
which there was a definite loss of nutrients.
of nutrients was accompanied

This loss

by a lowering of the con­

centration of the nutrients in the soil extract.

Later

in the growth the nutrients were again absorbed by the

-

4-

piants when the water extraotable nutrients were in­
creasing.

The phosphorus

absorbed by the plant and

that in the water extract did not decrease during this
period.

This is in agreement with the results obtained

by True and Bartlett (9) which showed "a definite con­
centration for each salt or mixture of salts at which
roots of peas absorb and excrete electrolytes at the
same rate.

If the culture solution is initially less

concentrated than this equilibrium concentration,
excretion from the roots over balance absorption.

If

the solution is initially more concentrated than this
equilibrium, absorption overbalances excretion".
Schreiner and Skinner (10) secured results which show
that "the higher the amount of any constituent present in
the solution, the more the plant growing in that solution
takes up this constituent, although it does not seem able
to use this additional amount economically.
Andre and DeMoussey (11) advanced the idea that the
difference in potassium and sodium content of plant tissues
is partially due to the uneoual mobility of the two alkali
metals in solution.
Much attention has been given to the influence of
fertilizer treatments upon the sugar content of beet
roots, but there is very little agreement on their effect.
Wiley (12) found a vast difference in the yield of beets
on various soils but the composition of the soils had

-

5-

little effect upon the sugar content of the beets*
Saillard (13) reported "that potassic fertilizers
gave good results in regards to purity, richness, and
yield of sugar beets.

Large applications of nitrate

of soda retarded maturity and decreased sugar content"*
Ifeibull (14) found "the sugar content was increased
slightly, but decidedly by nitrate of soda, while
phosphoric acid and potash

depressed the purity and

percentage of sugar in beets"*
Frankfurt (15) improved the yield of sugar content
of beets by the use of nitrate of soda and superphosphate
in combination.

Small applications of nitrate of soda

gave best results.
Vivier (16) concluded "that the beet crop increases
with applications of nitrate of soda and the density of
the juice decreases at about the same rate, so that the
total amount of sugar is not altered.

Phosphoric acid

shows little effect on the yield of crop and density of
juice"•
Joulie (17) reported that phosphoric acid and the
alkalies in the beet

roots varied with the richness in

sugar;-the phosphoric acid increased with an increase in
sugar, and the alkalies decreased under the same conditions*
IvIcOool and Harraer (18) found the sugar content of
beets to be greatly increased by the use of potash on
muck*

The increase was almost as great in beets from

-6-

plots fertilized with potash alone as in beets grown
on plots which received both phosphate and potash*
Schneidwind, Meyer, and Mflnter (19) increased
the dry matter and sugar content by the use of potash
on all soils investigated.

The best results were ob­

tained by fall applications.
Herke (20) reported that nitrate of soda applied in
moderate amounts in connection with potash and phosphoric
acid 6n soils poor in nitrogen increased the sugar content
of beets without injury to the quality.
The chemical constitution of enzymes is as yet a
matter of general speculation, so that investigators
have measured only the relative activity of enzymes in
preparations of plant tissues.

Bunzell (21) has used

the term "oxidase activity**rather than concentration of
oxidases to denote the factors responsible for the com­
plex reactions based upon oxygen absorption in the
presence of plant juices and tissues.

The decomposition

of hydrogen peroxide by plant and animal tissues was first
attributed to the action of a special enzyme by Loew (22)•
He termed it oatalase, and since that time it has been
demonstrated as present in all oxygen requiring cells.
Anaerobic cells contain no oatalase.

The only known

action of this enzyme according to Waldsohmidt (23) is
the decomposition of hydrogen peroxide into water and
oxygen.

-7-

There appears to "be a general agreement among
investigators that there is a correlation between health
and vigor in plants and the activity of oatalase in the
tissues.

Heincke (34) found less oatalase activity in

apple leaf tissuefrom trees grown on
in leaf tissue of

sandy soil than

trees grown on clay soil.

The cata-

lase activity was greater in leaves of trees grown in
cultivated soil than in leaves of trees grown in sod.
Applications of nitrate of soda to trees in sod increased
the oatalase activity of the leaves in proportion to the
amounts applied up to eight ounces per tree.
reported that nitrogen and phosphorus

Znott (25)

when applied to

the plant through the roots resulted in increased catalase activity.
Conversely a correlation has been found between
oxidase activity and retarded growth.
found the oxidase

activity to be much greater in sugar

beets affected with curly top
beets.

Bunzell (26)

disease than healthy

He found increased activity of oxidase in

every case where growth was retarded by natural or
artificial means.

Potato plants affected with curly

dwarf disease (27) gave a much higher oxidase activity
accompanied by greatly decreased growth as compared to
healthy potatoes.

Woods (28) reported a higher oxidase

activity in tobacco plants affected with mosiac disease
than in healthy plants.

-

8-

Ezell and Grist (29) on the other hand report^a
slight oorrelation between oxidase activity and growth
with a tendency to he negative in character.

The

correlation between oatalase activity and growth was
better and distinctly negative” .

The fertilizer

treatments described by Ezell and Grist are extremely
heavy and the absorption of these salts in excess may
explain this reversal of the activity of the enzymes.
Santesson (30) and Heincke (31) found that nitrates
of sodium, calcium, and potassium added to plant
tissues decreased their power to decompose hydrogen
peroxide•
In the investigations reported in this paper the
results of studies of the mineral nutrient content,
sugar content, and activities of the enzymes, catalase
and oxidase, of sugar beets as influenced by soil con­
ditions and fertilizer treatments are presented.
data

This

may contribute some knowledge to the problem of

the action of fertilizer salts and throw some light upon
the physiological role of nutrient elements.

Methods of Experimentation.
The

mineral nutrient content of leaves and roots

of sugar beets was determined on samples taken from

fertilizer plots on widely different soils.

Samples

were taken from plots on Eifle muck and Hillsdale
sandy loam at four periodp during the growing season
of 1926, and from Miami loam at four periods during
the season of 1927.

Samples of roots and tops were

taken from plots on Miami loam, Brookston silt loam,
and Berrien sandy loam at harvest time in 1927.
samples were placed in

These

moisture proof hags and taken

to the laboratory as quickly as possible.

Here they

were washed free from adhereing soil and dust, and
spread out to dry.

An electric fan was used to ac­

celerate the drying and prevent decomposition of the
material.

After the samples were thoroughly dried

they were ground in a Wiley Mill and preserved in glass
bottles for analysis.
Determinations of the mineral nutrients were made
according to the "Official Methods of the American
Agricultural Chemists" (32) with the exceptions of
potassium and sodium.

Determinations of potassium

were made by the cobaltinitrite method described by
Addie and Woods (33).

Sodium determinations were made

by the method described by Blancheterie (34) and
adapted to plant analysis by Bertrand and Perietzeanu
(35).

In this method the plant ash is dissolved

in

-10water and neutralized with acetic acid.

Phosphorus

,

the only element found in plant material which inter­
feres with the precipitation of sodium, is removed by
precipitation as uranium phosphate and centrifuging*
The sodium is then precipitated as the triple acetate
of uranium - magnesium - sodium*

The precipitate is

filtered through a weighed Gooch crucible, observing
the precautions of Blancheterie, washed with the rea­
gent and with alcohol, dried in the oven for one hour
at 110°C*, cooled in a dessicator, and weighed*

The

writer used Pyrex glass instead of the silica dishes
described by Blancheterie, and found that it was
necessary to run blanks with each set as there was an
appreciable amount of sodium in the glass which pre­
cipitated out*

When this amount was subtracted from

the weight of precipitate obtained by using 1 cc of
ff/lO NaOH the weight of sodium obtained checked very
closely with the weight of sodium present and did not
vary at any time more than *05 mg*

Since beginning

this work the writer has read Eoltoff’s description
of an improved method employing uranium - zinc acetate
instead of uranium - magnesium acetate for the
precipitation of sodium*

However the results obtained

by the method described above have been so satisfactory
that it was not considered desirable to change at this
time •

11-

Determinations of sugar content of sugar beets
was made on beet roots taken from fertilizer plots on
Rifle muck, Hillsdale sandy loam, Miami loam, Brookston
loam, Brookston silt loam, and Uappanee silt loam in
1926, and from Miami loam, Brookston loam, Berrien
sandy loam, and Nappanee silt loam in 1927*

These

samples were placed in moisture proof bags and trans­
ported to the College as quickly as possible, where
the sugar content was determined by the Experiment
Station Chemist.
Leaves of sugar beets were collected from the
fertilized plots on Rifle muck, Miami loam, and
Brookston silt loam for the study of the influence
of soil fertilizer treatments on the activity of the
enzymes^oatalase and oxidase.

These samples were

placed in quart mason jars, sealed, packed in ice,
and immediately taken to the laboratory.

The

preparation of the plant material was similar to
the method described by Ezell and Crist.
cut from the leaves with a cork punch.

Discs were
Two grams of

the discs were placed in a mortar with an equal weight
of pure calcium carbonate and sufficient distilled
water to cause the calcium carbonate to spread evenly
over the discs.

The material was ground with a pestle

until a uniform creamy misture was obtained.

The

sample was then transferred to a 100 cc. volumetric
flask and made up to

volume with water.

It was

-

12-

then preserved in a refrigerator at about 4° C until
used.

This is the temperature reported by Enott (25)

as being most favorable for the preservation of plant
material to be used for measuring the activity of
enzymes.

The determinations were made according to

the methods described by Ezell and Crist, except that
the mercury was removed from the manometer of the
Bunzell apparatus and the tube connected to- a mercury
displacement apparatus made from small graduated
pipettes.

The volume of oxygen split from hydrogen

peroxide was determined at 5, 10, and 15 minutes.

At

15 minutes this reading was usually constant and this
amount was taken as the measure of catalase activity.
The volume of oxygen absorbed by pyrogallol in one
hour in the presence of the plant material was ac­
cepted as the measure of the oxidase activity.

One

cubic centimeter of material was used for determining
catalase activity and three cubic centimeters for
oxidase activity.

The activity of each enzyme has

been calculated to cubic centimeters of oxygen per
gram of fresh plant material.

EXPERIMENTAL RESULTS
The Chemical Composition of Sugar Beets
The study of the chemical composition of the sugar
beets was divided into the

determination of the in­

fluence of season and fertilizer treatments on the

-13
mineral nutrient content of the leaves and roots of

-

beets grown on different soils, and the influence of
fertilizer treatments on the sugar content of the beet
roots grown on several soils*
I.
Seasonal Variation of Mineral Nutrients
The seasonal variations of the mineral nutrient
content of the leaves of beets grown on Rifle muck
and on Hillsdale sandy loam were determined for four
periods of growth during the season of 1926, and
similarly for beets grown on Miami loam for 1927.
These data are presented in Tables I, IX, and

III*

Table I presents a summary of the seasonal var­
iations of the mineral nutrient content of leaves of
beets grown on Rifle muck in 1926*
phosphorus

The percentage of

was high in the spring at thinning time,

but decreased rapidly during the summer, and following
this increased rapidly in the later stages of growth*
The highest percentage v/as found in the beets grown
on the plot which was fertilized with superphosphate
alone and was slightly lower in beets which received
potash in addition.

The percentage of potassium was

highest in the spring and shows a marked decrease during
the successive stages of growth.

An interesting

phenomenon noted at this point and which will be dis­
cussed later is that there was less potassium than

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Superphosphate (20$)

Seasonal Variations of Mineral Nutrient Content of Leaves of Sugar Beets grown on Rifle Muck

O

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-14phosphorus

in the leaves of beets from plots re­

ceiving no potash fertilization and that the potassium
content was greater than the phosphorus

content in

leaves of beets from the plots which were fertilized
with potash*

The percentage of sodium was high during

the early stages of growth and was higher in plants
from plots which received no potash than in those from
the potash treated plots*

During the later stages of

growth there was a decrease in the sodium content of
plants from all plots*

At this time the differences

in percentages of sodium in leaves of beets from the
different plots was slight.

The percentages of calcium

and magnesium were about equal during the first period
of growth.

The percentage of calcium decreased during the

growing season and increased at harvest time.

The per­

centage of magnesium followed a similar path in the beet
leaves from the plot which received both potash and
superphosphate.

The percentage of magnesium increased in

the leaves of the beets grown on the other plots during
the season and decreased at harvest time.
The largest beets were produced on the plot receiving
potash and superphosphate in combination.

It is interest­

ing to note the extremely high percentage of potassium,
followed by a high percentage of phosphorus

in the leaves

from this plot, and that the percentage of calcium was
greater

than that of magnesium.

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15-

The results abtained from beets grown on the
Hillsdale sandy loam are given in Table II.

Ac­

cording to these data the percentage of calcium
was extremely high in the leaves during the early
periods of growth and decreased rapidly during the
entire growing season.

The percentage of calcium

was highest in the leaves of beets produced on the
plot which received lime without potash.
phosphorus

The

content was low during the first period

of growth, increased in August, decreased in September,
and then increased in October.
was the phosphorus

An exception to this

in the leaves from the plot which

was limed,but received no potash. In these the percent­
age of phosphorous was high in the early stages of
growth, showed a slight decrease during the summer, and
then increased in the fall.

The percentage of potassium

was lowest in the leaves of the beets from all plots in
early July.

The percentage of potassium increased in

August, following which the beets grown on the plot
fertilized with urea and phosphate were the only ones
in which the percentage of potassium increased up to
harvest time.

A decrease in the percentage of potassium

was found in the leaves sampled in September, but no
further change occurred in October.

The beets from the

plot fertilized with urea, phosphate and potash, and
limed showed an increase at harvest time.

The magnesium

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Table III
Seasonal Variations of Mineral Nutrient Contetn of Leaves of Sugar Beets- grown on Miami loam - 19£7

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16content was highest at thinning time, after which there
was a gradual decrease, except in September, when there
was a slight increase, followed by a decrease at harvest
time.

The percentage of sodium was lowest in the cases

in which potassium was highest, and highest when the
potassium was

lowest.

Here again it is noted that the percentage of
potassium was greater than that of phosphorus

and that

of calcium greater than that of magnesium in the beet
leaves from the limed plot fertilized with urea, phosphate,
and potash, and that this plot produced the largest beets
grown on this soil.

The same is true of the plot which

was similarly fertilized but which was unlimed.

In the

beet leaves from the other plots the percentages of
phosphorus

were greater than the percentages of potassium.

The seasonal variations in the mineral nutrients in
the leaves of beets grown on Miami loam are presented in
Table III.
phorus

These data show a high percentage of phos­

in the leaves at thinning time, a decrease during

the summer, and then a large increase in the fall, except
in the case of beets grown on plots which received no
phosphate fertilizer.

The beet leaves on these plots

contained a low percentage of phosphorus

at thinning

time, but it gradually increased during the summer, and
was highest at harvest time.

The percentage of phosphorus

in the leaves from the phosphate treated plots was high at
thinning time, decreased during the summer, and then

-17inoreased during the later stages of growth*

The

percentage of calcium was lowest at thinning time and
increased gradually during the season except in the
beets from the non-potash treated plots*

In these oases

there was a decrease in the percentage of calcium in the
leaves at harvest time.

The percentage of magnesium was

low during the early periods, increased during the summer,
and then decreased at harvest time.

The percentage of

sodium in the leaves was low at thinning time and in­
creased during July and August, except in beets grown
on the plot with nitrate of soda,dnd superphosphate.

The

percentage of sodium decreased in the late stages of
growth except in the beet leaves grown on the plot fertil­
ized with nitrate of soda and potash.

The leaves from the

nitrate-phosphate plot showed a decrease in the percentage
of sodium during the entire growing season, while there
was an increase in the sodium content during the season
in leaves from the nitrate-potash plot.
Here also we find a high percentage of potassium
and a slightly lower percentage of phosphorus,

in the

beet leaves associated with the optimum growth of the
beet.

The percentage of calcium in the leaves of the

largest beets on the Miami loam was greater than the
percentage of m^n^esium except in the case of the July
samples.

It should be noted also that the percentage

of phosphorus

in the leaves of the largest beets is

greater than the percentage of calcium.

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-18-

II.
Variations in Mineral Nutrient Content of Beets
at Harvest Time Due to Fertilizer Treatments.
A summary of the results of the investigations
concerning the influence of fertilizer treatments and
soil conditions on the percentage of mineral nutrients
contained in the leaves and roots of beets are given in
Tables IV, V, VI, VII, and VIII.
Table IV presents the data obtained from beets
grown on Hillsdale sandy loam.

The use of potash,

especially muriate of potash, and of lime, either
singly or in combination, on plots supplied with nitro­
gen and phosphate increased the

yield of beets.

On the

unlimed plots the addition of potash increased the per­
centage of potassium in both leaves and roots, at the same
time causing a decrease in the percentages of calcium,
magnesium, sodium and phosphorus.
occurred in the

limed series.

The same phenomena

Lime in addition to urea

and phosphate greatly decreased the percentage of potassium
in the leaves and roots, but caused an increase in the per­
centages of phosphorus, calcium, and magnesium.

The leaves

of beets from the plots which produced the largest yield
of beets contained the minerals in the following order
P> Ca> Mg> Na, while in the roots the order was K > P > Na>
Ca> Mg.

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-19-

Table V. presents the data concerning the influence
of fertilizer treatments on the percentages of mineral
nutrients found in the leaves and roots of beets grown
on Rifle muck and sampled at harvest time.

The percent­

ages of phosphorus

and of potassium were extremely low

in the check plot.

Addition of potash caused a remark­

able increase in the potassium content, and a slight in­
crease in the phosphorus
roots.

content of both leaves and

Phosphate fertilization on the other hand caused

a tremendous increase in the phosphorus
slight increase in the potassium content.

content and a
Application

of 300 pounds each of superphosphate and potash caused
a large increase in production of both leaves and roots.
The percentage of potassium was slightly less than in the
beets from the plot fertilized with potash alone, and the
percentage of phosphorus

was considerably smaller than

in the beets grown on the plot which received superphos­
phate alone.

The percentages of calcium, magnesium, and

sodium were highest in the beets

from the plots showing

the smallest growth and least in beets from the plot
showing the maximum growth of both tops and roots.

There

was less variation in calcium and magnesium in roots
than in leaves and greater variation in sodium.
The results of the experiment on ^lami loam are given
in Table VI.

The application of^Ghilean Nitrate'4and

potash in combination, caused an increase in the percent­
ages of phosphorus , calcium, magnesium, and sodium and a

1927

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-20deorease in the percentage of potassium in the leaves.
In the roots the percentages of potassium and sodium
were increased, calcium and magnesium decreased, and
phosphorus

was unchanged.

Additions of 200, 400, and

800 pounds of superphosphate with the Chilean Nitrate
and potash caused corresponding increases in phosphorus
content and decreases in calcium and magnesium in both
roots and leaves with each succesive increment.

The

percentage of potassium was decreased in the roots with
successive addition of superphosphate.

The greatest

amount of potassium was found in the leaves from the plot
receiving 400 pounds of superphosphate.

Application of

800 pounds of superphosphate caused a decrease in the
amount of potassium in the leaves.

The percentage of

sodium was low in leaves from plots receiving 200 and 400
pounds of superphosphate but increased in leaves from the
plot receiving 800 po&nds.
The percentage of potassium was lower in the leaves
and roots of beets from the plots receiving nitrate of
soda and superphosphate than in the beets from the check
plot.

Additions of 50 and 100 pounds of potash caused an

increase in the percentages of potassium in the roots and
leaves and of phosphorus

in the leaves.

200 pounds of

potash caused a dedrease in both potassium and phosphorus
in leaves and in potassium in the root.
ment had no effect on phosphorus

The above treat­

content of the roots.

-21The percentages of calcium and magnesium in the leaves
were decreased by addition of 50 pounds of potash, in­
creased with application of 100 pounds, and then the
200 pound application caused a further increase in
magnesium content and a decrease in calcium content*
The percentages of calcium and magnesium

were slightly

decreased in the roots by additions of potash.

The

percentage of sodium was highest in the plot receiving
nitrate of soda and superphosphate only and decreased
with additions of potash.
fertilization with superphosphate and potash only
caused an increase in both potassium and phosphorus

con­

tent and a decrease in the calcium, magnesium, and sodium
in the leaves.

In the roots this

was accompanied by a

decrease in the percentages of all the elements except
sodium.

Additions of 100 and 300 pounds of "Chilean

Nitrate” caused an increase in the amount of phosphor us
and

sodium in the leaves and roots and a decrease in the

percentages of calcium and magnesium.

The potassium con­

tent was increased by the addition of 100 pounds of
"Chilean Nitrate”, but the percentage present in both leaves
and roots was decreased by the addition of 300 pounds.
The order in which the percentages of the elements
occurred in the leaves from the plot, which gave the
maximum yield of beets, was Z > P > N a > Ca> Mg and in the
roots P > £ > M g 3 N a > Ca.
The results of the experiment on Berrien sandy loam

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Table VII
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-22are given in Table VII.

The addition of 200, 400, and

800 pounds of superphosphate increased the percentages
of phosphorus,

in both leaves and roots and caused a

decrease in the percentages of calcium, magnesium, and
sodium as compared to the amount present in the plot
receiving "Chilean Nitrate" and potash only#

This

latter treatment caused an increase in the percentages
of sodium and phosohorus
calcium, and magnesium.

and a decrease in potassium,
The addition of 200 and 400

pounds of superphosphate caused an increase in the
potassium content of the leaves, while the 800 pound
application caused a decrease.

In the roots both the

400 pound and the 800 pound treatments caused a de­
crease in the percentage of potassium in leaves.

The

percentages of calcium and magnesium were lowest in the
leaves and roots from the plot fertilized with 400
pounds of superphosphate.
"Chilean Nitrate" and superphosphate alone increased
the percentages of phosphorus

and sodium in the leaves and

roots, and decreased the potassium, calcium, and magnesium#
Additions of potash up to 100 pounds per acre increased the
potassium in the leaves.

200 pounds of potash per acre

decreased the percentage of potassium in both leaves and
roots and caused an increase in the percentages of calcium,
magnesium,

[phosphorus > and sodium as compared to the plot

receiving only 100 pounds of potash.

The percentages of

calcium and magnesium present were low in this plot also.

of
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None

Fertilizer

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CP
CM
•
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CM
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to

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tf
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*
CO

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ID
O
Qt

60
IS
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CS
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—* * * * • •

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•
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to

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rH
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CM

Q
O
CM
rH

P 400
N 100
K
N 300
K

Leaves

•

IO
IO

O
O
CO
rH

N 100 P 4 0 0
K 90
N 100 P 4 0 0
K 200
N 100 P 4 0 0
K 20 0

•a

•

•

P 400

R o o ts

©
* •

•

N 100

Leaves

• •

0
01
* Oi*

N 100 P 8 0 0
K 100

R o o ts

N 100 P 4 0 0
K 100

0
23
6
"OC1
Q
7?
Ph

K100

Leaves

N100

O

Treatments

on Broolcston
grown
Beets
Sugar
of

R o o ts

Content
N u trie n t
M ineral

Table Till

HI

N 100 P 200
K 100

s ilt

loam

and

the

average

Q
3*
*
rH

-23Superphosphate and potash alone increased the
phosphorus content of leaves and roots but caused a decrease
in the percentage of potassium, sodium, calcium, and
magnesium.

100 and. 300 pounds of "Chilean Nitrate” per

acre caused a marked increase in the potassium and sodium
in the leaves, and a decrease in the percentages of
calcium and magnesium in both leaves and roots, and of
phosphorus
a

in the roots.

100 pounds of nitrate caused

decrease in the percentage of phosphorus

and 300

pounds caused a large increase in the leaves.
The percentages of elements in the leaves from the
plot which received 100 pounds of "Chilean Nitrate” , 400
pounds of superphosphate, and 100 pounds of potash occurred
in the following order K > N a > ,P’> C a > M g and in the roots
K>P>Mg>Na>Ca.
Table VTII gives a summary of the data obtained from
beets grown on Brookston silt loam.

The application of

"Chilean Nitrate” and potash in combination caused a decrease
in the percentages of potassium and magnesium in the leaves
and an increase in the phosphorus

calcium, and sodium,

while the percentages of potassium, magnesium, calcium, and
sodium were increased in the roots and the percentage of
phosphorus

decreased.

Additions of 200, 400 and 800 pounds

of superphosphate per acre caused marked., increases of
phosphorus ' in the leaves and roots and decreases in the
percentages of ealcium, magnesium, and potassium, with the
exception of the 800 pound application which caused an

-24increase in the potassium in the leaves.

The percentage

of sodium was decreased by the addition of 200 and 400
pounds of superphosphate and then increased by the 800
pound application.
The percentages of potassium, calcium, and magnesium,
in the leaves were reduced by the application of "Chilean
Nitrate" and superphosphate
and phosphorus

increased.

and the percentages of sodium
The sodium, phosphorus , and

potassium contents were increased in the roots by this
same treatment and the percentages of calcium and magnesium
lowered.

Additions of 50 pounds of potash increased the

content of the five elements in the leaves and of potassium
and sodium in the roots.

The 100 pound application caused

a decrease in the percentage of potassium, sodium, and
phosphorus1, and an increase in the calcium and magnesium
content in both leaves and roots, while the 200 pound
addition caused the potassium and sodium content to in­
crease, the percentages of calcium and magnesium to de­
crease, and had no influence on the percentage of
phosphorus .
Leaves from the plot fertilized with superphosphate
and potash show

a decrease in the percentages of

potassium, calcium, and magnesium, and an increase in the
amount of sodium and phosohorus *

The same treatment

caused a marked increase in the phosphorus
contents of the roots.

and potassium

Additions of 100 pounds of "Chilean

Nitrate" per acre caused a decrease in the percentages of

-25potassium and sodium in both leaves and roots, an in­
crease in the calcium, magnesium, and phosphorus

con­

tents of the leaves, and a decrease in the phosphorus #
calcium, and magnesium in the roots*

The addition of

300 pounds of "Chilean Uitrate" per acre caused a
further decrease in the percentages of potassium in
leaves and roots, a marked decrease
of phosphorus-

in the percentage

in the leaves, an increase in sodium

in both leaves and roots and of calcium and magnesium
in the leaves.

There was a slight decrease in percentages

of calcium and magnesium in roots.
The ratio of the elements in the leaves from the plot
producing the largest beets was K > P>Ca>'Ua7>Mg and of the
roots K > PP>Mg>-0a>ITa.

III.
Variations in Sugar Content of Hoots Due to
fertilizer Treatments on Several Soil
Types
The sugar contents of beets grown on several soils
under different fertilizer treatments in 1926 and 1927
are

given in Tables IX, X, XI, XII, XIII, XIV, and XV-.

A study of these tables

shows a very slight and insig­

nificant correlation between the fertilizer treatments
and the percentage of sugar found in the beets.

In

some cases there is a slight decrease in the sugar content

Table IX
In f l u e n c e o f v a r y in g am ounts o f " C h ile a n N i t r a t e " on S u g a r C o n te n t
o f B e e ts Grown on M iam i lo a m , B ro o kB to n lo a m , and B ro o k s to n s i l t lo am *
19 2 6
F e r tiliz e r
T r e a tm e n t
None

M ia m i
Loam

B ro o k s to n
Loam

B ro o k s to n
S i l t Loam

1 4 *5

1 7 .4

1 8 .4

1 6 .2

1 7 .4

1 7 .9

1 5 .1

1 8 .0

1 7 .4

1 5 .4

1 8 .3

1 7 .5

1 5 .7

1 7 .9

1 5 .7

1 5 .6

1 7 .5

1 6 .9

1 4 .7

1 8 .4

1 7 .6

4 0 0 # 0 - 1 2 - 6 N1Q0#

1 6 .4

1 6 .9

1 7 .8

400# 0 -1 2 -6 N200#

1 5 .7

1 7 .2

1 8 .1

400# 0 -1 2 -6

1 6 .5

1 6 .3

1 7 .1

4 0 0 # 0 - 1 2 - 6 N 300 #

1 6 .5

1 7 .4

1 7 .5

4 0 0 # 0 - 1 2 - 6 N 100 #

1 5 .8

1 7 .7

1 8 .3

400# 0 -1 2 -6

1 6 .0

1 8 .1

1 7 .8

4 0 0 # 0 - 1 2 - 6 N200#

1 6 .4

1 7 .5

1 7 .4

4 0 0 # 0 - 1 2 - 6 N 300 #

1 5 .0

1 7 .6

1 6 .5

400# 0 -1 2 -6

1 6 .4

1 7 .0

1 8 .5

4 0 0 # 0 - 1 2 - 6 N 100 #
None
4 0 0 # 0 - 1 2 - 6 N 200#
None
4 0 0 # 0 - 1 2 - 6 N300#
None

N - C h ile a n N i t r a t e o f Soda

0-12-6 Composed o f s u p e rp h o s p h a te and M u r ia t e o f P o ta s h

Table X
I n f l u e n c e o f Form s o f P o ta s h on S u g a r C o n te n t o f B e e ts grown on
H i l l s d a l e Sandy lo a m , M ia m i lo a m , and B ro o k s to n loam *
1926

F e r t i l i z e r T re a tm e n t

M iam i
Loam

B ro o k s to n

H i l l s d a l e Sandy
loam *

Loam
U n liraed

U 50 P 1 0 0 c k

1 5 .6

1 6 .1

1 6 *3

1 6 .8
1 6 *7

"

"

KCL 1 0 0

1 5 *7

1 7 .7

1 7 .1

"

"

ck

1 6 .0

1 6 .1

—

"

n K SO 1 0 0
2 4

1 6 .6

1 6 .9

1 6 .0

1 6 .1

**

»

1 4 *2

1 6 .3

1 5 .9

1 6 .4

"

"

15 *9

1 7 .5

1 6 .0

1 6 .6

1 5 .8

--

--

---

1 5 .2

1 7 .3

1 6 .4

1 6 .0

1 6 .4

1 6 .6

—

---

1 5 .2

1 8 .6

--

--

1 6 *5

1 7 .7

--

--

1 6 .9

1 7 .3

ck
ECL 100

None
U 50 P 1 0 0

I S O 10 0
2 4

None
U 50 P 1 0 0

KCL 100

None
U 50 P 10 0

K So 1 0 0
2
4

—

— —

1 7 .3

None

U

Lim ed

• U re a

P = A naconda 4 5 $
* On

H i l l s d a l e Sandy lo am u re a was u sed a t r a t e

CAH (PO \

4

*'2

o f 3 3 # p e r a c r e and

a t 7 1 # p e r a c r e i n p la c e o f U 50 P 1 0 0 *

Table XI
Influence of Time of Applying "Chilean Nitrate" on Sugar
Content of Sugar Beets on Brookston silt loam
1926

400# 0-12-6 ck

18.0

No Fertilizer

17.3

it

IN 150#

16.5

400# 0-12-6 3N 50#

16.5

»

SN 75#

17.9

"

16.0

4N 38#

w

ck

18.1

No Fertilizer

17.1

»

3N 50#

17.9

400# 0-12-6 In 150#

16.3

tt

4N 38#

16.4

i»

ck

17.7

No Fertilizer

16.6

17.2

400# 0-12-6 3N 50#

17.3

tt

IN 150#

tt

2N 75#

18.0

tt

ck

17.8

"

"

2N 75#

17.5

4N 38#

17.1
18.0

Fertilizer

IN 150 - 1 application of 150# Chilean Nitrate of Soda May 7
2N 75 :

2

n

each of 75# "

*

"

" May 7, & June 2

3N 50

m 3 applications each of 50# Chilean Nitrate of Soda May 7,
June 2 and 17.

4N 38

s 4 applications each of 38# Chilean Nitrate of Soda May 7,
June2, 17 and 29.

0-12-6 composed of superphosphate and muriate of potash.

Table XII
Influence of Varying Amounts of Superphosphate and potaslfc on Sugar
Content of Beets Grown on Nappanee silt loam*
1926
Phosphate
Treatments

Potash
Treatments

Ko

K 50

K 100

K 200

N 100 Po

13*0

18.5

18.6

16.8

N 100 P 200

17.4

18.5

19.2

17.3

N 100 P 400

17.1

18.5

18.7

16.5

N 100 P 600

18.6

17.4

17.6

N 100 P 100

20.5

18.2

18.2

No Fertilizer

18.0

—

N

• Chilean Nitrate of Soda

P a Superphosphate (20$)
K m Muriate of Potash

16.3
19.0

TABLE XIII
Influence of Varying Amounts of "Chilean Nitrate", Super­
phosphate, and Potash on Sugar Content of Beets in 1927*
Brookston
silt loam
Tuscola
County

Treatment

Miami
Loam
Eaton
County

Miami
Loam
Clinton
County

Berrien Sandy
Loam
Bay
County

No Fertilizer

17.2

17.2

15.8

17.2

No P40Q KLOO

17.1

17.3

16.8

18.6

N50 P400 K100

17.0

18.7

16.3

18.6

N100 P40Q KLOO

17.5

17.3

16.4

18.4

N200 P4QQ KlOO

16.9

17.4

15.4

17.3

N300 P400 KLOO

15.8

17.3

14,4

17.4

N100 PO KLOO

16.0

18.1

15.5

17.4

NlOO P200 KLOO

17.6

19.0

16.4

17.4

N100 P400 KlOO

17.5

17.4

17.3

18.4

NlOO P600 KlOO

16.5

18.3

16.9

NlOO P800 KlOO

17.4

18.1

16.1

18.7

NlOO P400 KO

17.1

17.0

16.1

18.4

18.2

"

K25

17.3

17.7

16.4

18.8

"

K50

16.6

17.5

16.4

18.8

*

KlOO

17.5

17.4

16.9

18.4

«

K200

17.3

17.9

15.7

18.4

N 100 s 100 pounds "Chilean Nitrate."
P 400 s 400 pounds of 20$ Superphosphate
K 50

s 50 pounds of Muriate of Potash

Table XIV
I n f l u e n c e o f V a r y in g C o m b in a tio n s o f " C h ile a n N i t r a t e , "
S u p e rp h o s p h a te and P o ta s h on th e Sugar C o n te n t o f B e e ts in
1927.
F e r tiliz e r
T re a tm e n t

B ro o k s to n
s i l t loam

Nappanee
s i l t loam

None

1 8 .1

1 7 .0

NO P 200 K50

1 8 .0

1 7 .9

N50 P 200 K50

1 8 .1

1 7 .7

NlOO P 200 K50

1 7 .8

1 6 .9

N 200 P 200 K50

1 6 .2

1 6 .9

None

1 7 .9

1 6 .7

NlOO P0 K50

1 7 .4

1 8 .1

NlOO P0 KlOO

1 7 .3

1 8 .1

NlOO P 200 K0

1 7 .8

1 7 .4

NlOO P 200 K50

1 7 .8

1 6 .9

NlOO P 200 KlOO

1 7 .7

1 6 .9

NlOO P 400 KO

1 7 .1

1 7 .4

NlOO P 400 K50

1 6 .3

1 6 .9

NlOO P400 KlOO

1 6 .9

1 7 .3

NlOO P800 KO

1 7 .0

1 7 .1

NlOO PQOO K50

1 6 .4

1 7 .7

NlOO P 800 KLOO

1 7 .7

1 8 .2

None

1 8 .0

1 6 .8

N - Chilean Nitrate of Soda
P - Superphosphate (20$)
K - M u r ia t e o f P o ta s h

-26of beets grown in the presence of extremely large amounts
of nitrate.

The plots which received delayed applications

of nitrate on Brookston silt loam showed a lowered sugar
content also.
The sugar content of beets grown on Rifle muck was
greatly increased by the use of potash.

The plot re­

ceiving 300 pounds each of potash and superphosphate
produced the largestyield of beets
of sugar was highest

in these.

and the percentage

The results on mineral

soils however are not significant.
It appears that the soil itself has a greater in­
fluence on the sugar content of the beets than does the
fertilizer treatments, which is probably due largely to
the moisture conditions.
The Influence of Fertilizer Treatment
on the Activities of Oxidase and
Gatalase.
In this experiment
sized plants growing

leaves wereselected from average

onfertilized plots on Rifle muck,

Miami loam, and Brookston silt loam during the season of
1927.

The beets on Rifle muck were

mounts of superphosphate and potash.

grown with varying

a-

The plot receiving

300 pounds of superphosphate and 300 pounds of potash
produced the maximum growth of tops

and roots. The

catalase activity as shown in Table

XVI was greatest

and the oxidase activity least in leaves of beets grown

TABLE XVI

'Enzyme Activity of Sugar Beet Leaves

Rifle Muck

Gatalase Activity
Treatment

Oxygen Evolved From R 2°2 *7
1 Gram Fresh Beet Ieaf
June 27

Aug. 13

Sept. 30

No Fertilizer

190 cc

150 cc

2G0 cc

£300

215 ”

175 "

230 "

£300 P100

230 ”

190 tt

230 n

£300 P300

270 "

225 "

325 »

£100 P300

270 "

220 "

300 "

P300

255 "

225 «

305 »

Oxidase Activity
Treatment

Oxygen Absorbed by Pyrogallol
In Presence of 1 Gram Fresh
Beat Leaf
June 27

l.ug. 13

Sept. 30

8.3 cc

8.0 cc

K300

4,0 ’»

5.0 «

3.8 "

£300 P100

3.5 "

4.0 "

3.0 "

£300 P300

1.7 "

1.7 "

2.0 "

£100 P300

4.0 "

5.0 "

4.5 »

5.0 "

6.4 "

P300

LO

7.5 cc

o•

Ho Fertilizer

-27-

on this plot at all three samplings.

Additions of

superphosphate appeared to hare a marked influence on
the activity of catalase as plots receiving 300 pounds
of superphosphate exhibited a very high catalase
activity, even though the application had no effect on
the growth of the plant as in the case of the plot
receiving superphosphate alone, since here the growth
was about the same as that on the check plot#

With

this exception the activities of catalase and oxidase
are

quite closely correlated with vigor of growth.

Beet leaves from fertilizer plots which showed increased
growth exhibited high catalase activity and a low oxidase
activity, the greatest catalase activity and the least
oxidase activity occurred in leaves of beets from the plot
producing the largest yield.

The least catalase activity

and the greatest oxidase activity occurred in leaves of
beets grown on the plot receiving no fertilizer in which
growth was very small.

The activity of catalase was

greatest in samples selected September 30 as compared to
samples selected in June and August.

The fall rains

produced an increased vigor of growth which probably
accounts for this increased activity of catalase.

The

activity of oxidase was not changed much, but remained
more constant for the three samplings.
The plots on Miami loam were fertilized with mixtures
of "Chilean Nitrate", superphosphate, and potash.

Maximum

yields of beets were secured on plots receiving 100 pounds
Chilean Nitrate, 400 pounds superphosphate, and 50 pounds
potash, although the largest leaves were produced on the.

TABLE XVII

Enzyme Activity of Sugar Beet Leaves

Miami Loam

Treatment

Catalase Activity______
Oxygen Evolved From
1
H 2 O2 By 1 Gram Fresh
Beet Leaves Aug, 1

Oxidase Activity
Oxygen Absorbed by Pyrogallol in Presence of 1
gram Fresh Beet Leaves

No Fertilizer

125 cc

10,0

NlOO KlOO

150 "

8.5

NlOO P200 KlOO

200 "

6.0

NlOO P800 KlOO

250 "

2.5

NlOO P400

310 n

1.7

NlOO P400 X50

330 H

1.7

NlOO P400 K200

320 "

3.5

P400 KlOO

210 "

2.0

N50 P400 KlOO

310 ”

1.6

N3Q0 P400 KlOO

325 "

1.6

FT = Chilean Nitrate of Soda
P - Superphosphate 20%
K = Muriate of Potash

-28plot receiving 300 pounds Chilean Nitrate in combin­
ation with superphosphate and potash.

The smallest growth

of tops and roots occurred on the plot receiving no fer­
tilizer and on the one receiving nitrate and potash only.
The extremely heavy applications of phosphate and potash
retarded the growth of the beets.

The data given in

Table XVTI shows the oxidase and catalase activities.
The highest catalase activity was exhibited by the
leaves of beets grown with 100 pounds Chilean Nitrate,
400 pounds Superphosphate, and 50 pounds potash.

The

lowest oxidase activity was produced in the leaves of
beets grown on the plots fertilized with 50, 100 and
300 pounds of Chilean Nitrate in combination with 400
pounds superphosphate and either 50 or 100 pounds of
potash.

The leaf growth on these plots was all about

the same.

The lowest catalase and the highest

oxidase

activity was produced by leaves of beets grown on the
check plot and the plot receiving Chilean Nitrate and
potash only.

On these plots an increased catalase

activity and decreased oxidase activity are associated
with

more vigorous growth, and decreased catalase

activity and increased oxidase activity with a retarding
of growth.
The beets grown on Brookston silt loam were fertil­
ized with varying amounts of "Chilean Nitrate", super­
phosphate, and potash.

The plot receiving 100 pounds

"Chilean Nitrate", 200 pounds superphosphate, and 100
pounds potash produced the largest yield of beets,

T a b le X V I I I
Enzyme A c t i v i t y

of

S ugar B e e t L eaves

B ro o k s to n s i l t Loam
C a ta la s A c t i v i t y

T r e a tm e n t

Oxygen e v o lv e d fro m
H „0g By 1 gram F re s h
Leaf
J u ly 2 2 , 1927

O xid ase A c t i v i t y
Oxygen A bsorbed by
P y r o g a llo l in P re ­
sense o f 1 gram
F re s h B e e t L e a f
J u ly 2 2 , 1927

No F e r t i l i z e r

40

1 0 .2

NO P 200 K5Q

65

1 0 .2

N 100 P 2 0 0 K50

12 5

6 ,8

N 100 P200 K100

15 0

3 ,0

N 100 P 400 KO

100

N 100 P400 KO

100

6 .8

N100 P 800 KO

14 0

3 .0

N 100 P 800 KLOO

140

3 .4

N -

C h ile a n N i t r a t e o f Soda

P -

S u p e rp h o s p h a te

{2 0 fo)

K - M u r ia t e o f P o ta s h

8 .5

-29closely followed by the two plots receiving 800 pounds of
superphosphate.

The smallest beets were produced on the

plot receiving no fertilizer and on the one receiving only
200 pounds superphosphate and 50 pounds potash.

The

activity of catalase and oxidase were here, too, quite
closely correlated with vigor of wrowth as shown in Table
XVIII.

The highest catalase activity was found in beet

leaves from plot receiving 100 pounds "Chilean Nitrate",
200 pounds superphosphate, and 100 pounds potash.

The

least catalase activity and the greatest oxidase activity
was exhibited by leaves of beets grown on the check plot.
A comparison of the three tables shows a decidedly
lower activity of catalase in beet leaves from the
Brookston silt loam than in leaves of beets from Miami
loam or Rifle muck, although the yield of beets was much
greater.

However, these samples were taken during a

period of great drought, while the samples from Miami
loam were taken after a rain.

The samples from Rifle

muck were secured at times when moisture conditions were
conducive

to good growth.

This probably accounts for

the difference in activity of catalase as the growth
would be less vigorous during a dry period.
Discussion
The data presented show a marked variation in the
mineral nutrient content, sugar content, and enzyme
activities of sugar beets grown under various soil con­
ditions and fertilizer treatments.

There is a marked

-30correlation between the mineral nutrient content of
beet leaves and

roots, activities of enzymes, and the soil

conditions and fertilizer treatments.
ficant correlation between these

There is no signi­

conditions and the sugar

content of the beets.
There is no definite seasonal absorption of the
mineral nutrients on the three soils studied similar to
the absorption described by Burd, but the percentages of
the various nutrients present at any one time appear to
be

dependent upon the soil itself, the fertilizer ap­

plied, and the stage of growth.

On Rifle muck the per­

centage of potassium in the plants was highest at the
beginning of growth and then decreased rapidly.

It

appears that the available supply of potassium was ab­
sorbed from this soil by the plants in the early stages
of growth and
growing season.

was then utilized during the rest of the
There was no accumulation of potassium

during the later stages of growth, so there must have
been less available potassium in this soil than was re­
quired by the plants, since Loew has shown that plants
require a minimum of each nutrient mineral and when this
is supplied they usually take up not only an excess of
these, but also

quantities of other compounds present

in the medium solution.

Hoagland also found a marked

absorption of nutrient elements at all stages of growth
when suitable concentrations of ions were maintained.

The percentage of potassium in the beet leaves grown on
Hillsdale sandy loam was lowest at the beginning of the
growing season, although the beets from the plot which
was limed, and fertilized with urea, phosphate and
potash were the only ones which showed an accumulation
in the fall*

On Miami loam in 1927 all plots except

the check showed an accumulation of potassium in the
leaves during the final stages of growth.

All of

the plots on all three soils showed an accumulation of
phosphorus,

in the leaves at harvest time.

On Rifle

muck calcium tended to decrease during the midseason and
increase in the fall, while on the other hand magnesium
increased during the summer and was much lower in the
fall.

Sodium like potassium was highest in the early

season and lowest at harvest.

The calcium and magnesium

contents were highest during the first:periods of growth
and lowest in the final stages of growth on Hillsdale sandy
loam.

The percentage of calcium in the leaves of beets

on Miami was lowest in the spring and tended to increase
up to the date of final sampling on all plots receiving
muriate of potash in the fertilizer.

On the other two

plots, there was a decrease in calcium at harvest.

The

magnesium increased during the summer, but decreased in the
fall.

It is quite evident from the above that the potash

fertilizer must have released calcium by base exchange.
It is evident also that the Seasonal fluctuations in
the mineral nutrient content of beets are influenced
largely by the two factors, concentration of the ions in
the soil solution and the stage of growth.

The concentration

-32of the soil solution is controlled by the nature of the
soil itself, the fertilizers applied, moisture conditions,
temperature•
It is shown in this work that

the mineral nutrient

contents of sugar beets, vary not only with the soils
upon which they are grown but also with the fertilizer
treatments afforded each soil.

The percentage of a

mineral nutrient present is also influenced by the
growth of the plant.
be traced

Fraps (36) states "no relation can

between the additions and the phosphate con­

tent of crops.

When the crops are unusually small the

phosphoric acid usually runs higher than the average".
Under the conditions of the experiments reported in this
work, however, it has been shown that the phosphorus
content of sugar beets was greatly increased by every
addition of superphosphate although the size of the
beet was doubled ill some cases.
■^udge (37) and Spurway (38) have both pointed out
that sodium nitrate exerts a beneficial effect in render­
ing soil phosphate available.

The applications of "Chilean

Nitrate" on Miami loam each caused a correspondingly great
increase in the phosphorus content of the roots and leaves
of beets.

On 3errien sandy loam the smaller application

caused a decrease in the percentage, while the larger
application caused an increase in the phosphorus content.
On Brookston silt loam the results were just the reverse.

-33It is evident therefore that nitrate of soda exerts a
remarkable influence in rendering the phosphate in the
soil more available to plants.
Potash salts added to a soil deficient in potassium
and with a low supply of available phosphorus cause a de­
crease in the phosphorus content of the beet plants.

This

is not due to the phosphorus becoming unavailable but to
the fact that the increased growth of plant utilizes the
amount of phosphorus more economically.

The smaller a-

mounts of potassium added in conjucntion with nitrate and
phosphate increased the phosphorus but the heavy applica­
tion reduced it except in the Berrien sand.

Here the 100

pound application decreased and the 200 pound application
increased the phosphorus content.
On Miami loam the fertilizer treatments which resulted
in increased growth caused an increase in the percentage of
potassium in the leaves of the sugar beets.

The excessive

applications of potash as well as of nitrate and super­
phosphate caused a decrease in the percentage of potassium
in the leaves, and resulted in a decrease in the growth of
the beet root.

The same is true in the case of beets

grown on Berrien sandy loam except in the comparison of
applications of "50 and 1 0 0 'pounds of potash.
On Brookston silt loam each fertilizer treatment which
resulted in increased growth caused a decrease in the
potassium content, and each treatment which resulted in a

-34decrease in growth gave an increase in potassium con­
tent of the leaves.
Fertilizers containing no sodium applied to beets,
such as phosphate and potassium caused a decrease in the
percentage of sodium in the plant.

When sodium was

applied however there was an increase in the sodium content.
Every successive increment of sodium nitrate caused a
like increase in the sodium content of the leaves,

lime,

potash and phosphate caused decreases in sodium in cases
where no sodium was applied in the fertilizer.
In most cases the percentages of calcium and mag­
nesium were lowest in the leaves and roots of the
largest beets.
The addition of varying amounts of the different
nutrient elements to the soil usually resulted in a
corresponding increase in the percentages of these ele­
ments found in the beet plants, except in the case of
additions of large amounts of potash.

Excessive appli­

cations of potash generally caused a decrease in the
percentage of potassium in both leaves and roots of
beets •
Additions of any one fertilizer constituent which
resulted in an increase in the growth of the beet plant
usually resulted in an increase in the percentage of this
element in the plant, and a decrease in the percentages of
other elements in it.

However as in the case of additions

of sodium nitrate which causes the phosphate to become

-35-

more readily available this is not true*
The fertilizer treatments in which the ratio of con­
stituents was most suitable for the production of the crop
resulted in an economical utilization of all the elements
needed in the growth of the plant*
The decrease in the growth of beets noted in most
cases in which excessive amounts of the various fertilizer
constituents were applied may oossibly be due to a too
high concentration of the soil solution.

According to

True and Bartlett (39) there is a definite concentration of
the. solution at which plants absorb and excrete ions at
the, same rate.

If the concentration of the solution is

greater than this there is an excretion of the ions from
the plant.

In case the concentration approached

this

equilibrium concentration it is likely that the plant
would not secure a sufficient supply of the nutrient.
The relative percentages of the elements as they
occur in the beets from plots which produced the largest
growth on the five soils investigated were found in the
following order K> P > C a > M g

in the leaves.

roots the order is £ > P > M g > C a .

In the

The relative amounts

in the leaves of beets from the plot on the Brookston
silt loam yielding the largest beets were 14: 7:2^5:1.5.*
On Hillsdale sandy loam the lowest yielding soil the
relative amounts were 8 :7:3:1.5 and on Kifle muck
8:6:2,.5;2.5.

There is evidently a definite ratio of the

-36mineral nutrients used by the plant which is optimum
for the production of that plant, and with sugar beets
studied the relative amounts given for beets grown on
Brookston silt loam seemed to be the best, as the beets
on this soil greatly out yielded those on all other
fields studied.
The

ratio of elements in beet leaves of the check

plot on Rifle muck was p) C a > K > M g ;

on Hillsdale sandy

loam P > K ^ > C a > M g although at the beginning of the
season it was Ca> Mg> P>K; on Berrien sandy loam K> >P Ca>
Mg, and on Brookston silt loam K > P > C a > M g .

It would

appear from this that toxicity might have some influence
in the small yield of beets on the soils and plots in
which the magnesium content of the leaves becomes greater
than the calcium content, and in which the potassiumcon­
tent becomes lower than the P or Ca content.
It has been quite generally conceded that phosphorus
is needed in the fertilization of the crop but that it has
little influence on the sugar content, while potassium has
acquired

a reputation for improving it, and sodium nitrate

for having a detrimental effect on it.

V/hile it is well

known that potassium is essential in the production of
sugar in the plant, there does not appear to be any
correlation between the fertilizer applied and sugar content
even in cases in which the potassium content is greatly
increased, exdept in the case of Rifle muck in which the

-37potassium content is extremely low*

The sugar content

was slightly decreased in some cases by heavy fertili­
zation with nitrate of soda, and not in others.

Addition

of phosphate to the fertilizer increased the sugar content
slightly but not significantly.

There is evidently little

danger of injury to the sugar content of beets under the
conditions of this experiment.
The hypothesis that

oxidase activity is associated

with decreased growth and as has been suggested in this
work and in the work of Bunzell (17)

(18), Woods (19)

and Ezell and Grist (20) seems to be correct.

In every

case reported (Table X) the oxidase activity was greatest
in the leaves of beets from those plots which produced
the smallest growth.

Likewise the hypothesis that

catalase activity is associated with increased growth as
pointed out by Heincke (15), also appears to be correct.
The data presented

in Table X shows the catalase

activity was greatest in the plots producing the greatest
growth.

In case of beets on Rifle muck the catalase

activity was associated with phosphate fertilization.

It

is quite evident that there was a positive correlation
between the catalase activity and vigor of growth of the
beets and a negative correlation of oxidase activity.
Similar results were obtained (unpublished data) by the
writer in studying the influence of light intensity on
growth and enzyme activities of sugar beets, in which

-38there was also a positive correlation between vigor of growth
and catalase activity and a negative correlation with oxidase
activity•

Summary

In these investigations the influence of soil con­
ditions and fertilizer treatments on the mineral nutrient
content at four periods of grov/th and at harvest time, on
the activities of the enzymes, oxidase and catalase, and
on the growth of sugar beets have been studied.

The re­

sults were as follows:1*

There are certain soils which are especially

adapted to the growth of sugar beets and others are un­
suitable for their production.

Brookston silt loam is

the best producer of sugar beets and Hillsdale sandy
loam, Berrien sandy loam, and Rifle muck lowest producers
naturally of sugar beets.
2.

Elements which are deficient in soils when

supplied in sufficient amounts cause the poor soils to
become fairly productive.
3.

Complete fertilizer plus lime gave highest yields

of beets on Hillsdale sandy loam.
4.

300 pounds each of superphosphate and potash

applied in combination gave largest beets on Rifle muck.

-39-

5.

A ratio of 100 pounds of Chilean Hitrate, 400

pounds of 20% superphosphate, and 100 pounds of potassium
muriate produced the largest beets on Miami loam, Berrien
sandy loam, and Brookston silt loam.
6.

The percentages of mineral nutrients in the

leaves of beets from above mentioned plots in the following
order K > P > C a > M g
7.

and in the roots K ^ P > M g > C a .

The ratio of the elements in the leaves, in the

order named, from the plot on Brookston silt loam yielding
the largest beets was 14:7:2.5:1.5.

On Hillsdale sandy

loam and Hifie muck, soils producing small roots, the
ratios were 8:7:3:1.5 and 8 :6 :2 .5:2.5 respectively.
8.

When the ratio of fertilizer elements applied is

such as to cause an increase in growth of the beet plant
there is a more economical utilization of the other
minerals taken from the soil.
9.

Applications of incomplete mixtures usually cause

an increase in the content of the elements applied and a
decrease in the percentages of other elements.
10.

Mixtures of two fertilizer salts (such as nitrate

and potash) sometimes cause decreases in the yield of beets as
in the case
11.

with Berrien sandy loam.

Boils and fertilizer treatments had very little

influence on

the

sugar content of beets except in the

case of Rifle muck.
12.

There is a positive correlation between cstalase

activity and vigor of growth in sugar befet leaves.
13.

There is a negative correlation between oxidase

activity and vigor of growth in sugar beet leaves.

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