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RED PINE (PINUS RESINOSA)
SEEDLINGS AS AFFECT?” BY
VARBOUS SOIL TREA'FMENTS

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thesis entitled

Red Pine (Pinus resinosa) Seedlings as

Affected by Various Soil Treatments.

presented bg

Orville Moore

has been accepted towards fulfillment

of the requirements for

Masters Soil Science

degree in

£272. 7:11»sz

Major professor

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RED PINE (PINUS RESINOSA) SEEDLINGS
AS AFFECTED BY VARIOUS SOIL TREATMENTS

By
Orville W. W

A THESIS
Submitted to the School of Graduate Studies of Michigan
State College of Agriculture and Applied Science

in partial fulfillment of the requirements

for the degree of

MASTER OF SCIENCE

Department of Soil Science

1950

ACKNOWLEDGEMENT

I wish to express my sincere gratitude to
Dr. Ray Cook for his help and guidance throughout
the experiment and in the preparation of this paper.
I am also indebted to the other faculty members and
students of the Soil Science Department who gave
helpful advice. I am especially grateful to Professor
Forrest Strong of the Botany Department for his interest
and suggestions, and for the seedlings which he

generously furnished for the experiment.

{uninfea

TABLE OF CONTENTS

INTRODUCTION

REVIEW OF LITERATURE
HISTORY OF THE SITE
EXPERIMENTAL PROCEDURE
DISCUSSION AND RESULTS
SUMMARY AND CCNCLUSIONS
TABLES

FIGURES

REFERENCES

Page

21
32
35
52
56

RED PINE (PINUS RESINOSA) SEEDLINGS
AS AFFECTED BY VARIOUS SOIL TREATMENTS

By
Orville W} Moore

INTRODUCTION

Early in the growing season of 19h9, the second
year seedbeds of red pine VEEEEE resinosa) and Norway
spruce [giggg‘ggig§), at the Michigan State College
Nursery, began to show signs of chlorosis. The
chlorosis gradually became more serious until by
late summer approximately 95 percent of the seed bed
area was affected, and, in some spots, seedlings had
died. The normal plants were distributed over the
area in small spots. Each spot constituted approxi-
mately one to four square feet.

Field observations and studies of the plants
failed to indicate the presence of disease organisms
or insects, so it was concluded that the problem
might be one of soil.

It was the purpose of this investigation to show
the cause of the stunted growth, chlorosis, and spotty

normal growth; also to show comparative effects of

various fertilizer and organic matter treatments by
measuring the growth response of red pine seedlings.

REVIEW OF LITERATURE

Clark (1916), by making total plant analyses,
determined that white pine (Pinus strobus) 2-0 seed-
lings, having an average density of 100 per square
foot, had removed 9t.6 pounds of available nitrogen,
31.8 pounds of phOSphoric acid (P205), and h1.6 pounds
of potash (K20) from an acre of soil. Retan (l9lh)
showed the beneficial effects of various inorganic
fertilizer treatments in combination with cOWpeas and
oats plowed under as green manure. Retan (1915) also
found that a charcoal application to heavy soils im-
proved aeration, drainage, and moisture retention, and
also helped to prevent "damping off". Wilde (l9h6)
shows that soil aeration data has the same significance
for all soil types. He states that watering should not
lower the aeration of the soil below 20 percent by vole
ume for any length of time to prevent denitrification
and other soil reduction processes. Toumey and Korstian
(1916) recommend the plowing under of legumes and the use
of heavy applications of farm manure in maintaining nurs-
ery fertility. Wilde (1937) stresses the importance of
organic matter in nursery soils, but he also states that

manure of any kind is undesirable, because of danger of

diseases and of spreading weed seeds. Wilde (1936) also

shows the danger from nematode infection and consequent
destruction of seedlings when barnyard manure is used.
Auten (l9h3) found that acid peat had a favorable
effect in correcting alkalinity and in improving den-
sity and height of pitch pine (ElEEE regida) and short-
leaf pine (2133; echinata) in forest nurseries of the
central states. Wilde and Hull (1937) recommend the
use of strongly acid peat as a fertilizer and as a
buffering material. These workers have shown that
strongly acid (pH 5.5 or lower) peat is valuable as a
base exchange material and as a nutrient amendment
particularly when its nitrogen content is 2 percent or
more. The beneficial effect of acid peat in coniferous
nursery soils is primarily the result of the change it
causes in soil reaction. It is also valuable in pro-
viding favorable soil structure. When nursery soils
are low in colloids, fertilizer salts are easily
leached and lost. When drought occurs, evaporation
causes the salts to rise to the surface with a conse-
quent "burning" of the roots of seedlings. Acid peat
'having a base exchange capacity of 80 or 100 m.e. per
100 grams is considered very satisfactory. All types

of peat are of minor significance so far as phosphorus

and potassium are concerned. However, their calcium,
sulphur, and iron contents exceed the minimum require-
ments of coniferous seedlings. The authors recommend
that applications he made so that the base exchange
value is brought to a desirable level. It should be
worked well into the upper 8 inches of the soil.
Eliason (1937) demonstrated that the color of buck-
wheat, when used as a cover crOp, is a fair indication
of the need of the soil for nitrogen. Lunt (1938)
reports injury to plants by the use of ammonium sulfate
on sandy soils of low buffer capacity. On such soils
he found concentrated nitrogen salts most effective
'when used in small doses at frequent intervals. Addi-
tion of calcium caused needle burning where phosphorus
and potassium.were low. Red pine was found to take up
less calcium than any other species tested. Lutz and
Chandler (19h6) point out that large amounts of calci-
um carbonate cause "damping off" and nutritional dis-
orders in conifers. Mitchell (l93h) shows that seed?
ling weight is influenced by seed weight. Root-shoot
ratio has been a standard index of seedling quality
(Wilde, 19h6; Mitchell, 193A; Toumey and Korstian,

1916). Mitchell found root-shoot ratios greater where

nitrogen was a limiting factor. Nitrogen and phos-
phorus deficiency color symptoms on foliage are simi-
lar to those of most agricultural crOps (Lutz & Chand-
ler, 19L6;‘Mitchell, 193k; Cook & Miller, l9h9).

Wilde (l9h6) states that green manures are most
valuable as "catch" craps for commercial fertilizers
by fixing a considerable portion as difficultly solu-
ble organic compounds. These are slowly made avail-
able over a period of time with little loss through
leaching.

Sandy loams and loamy sands are recommended tex-
tures for coniferous nursery soils according to Toumey
and Korstian (1916) and Wilde (l9h6). Wilde states
that the soil reaction range 5.0 to 6.0 is Optimum
for most conifers.

According to Toumey and Korstian (1916) coniferous
seedlings absorb nitrogen in greater amounts than they
do other elements. They believe the fertilizer ratio
used should be lO-h-S for best results. Concentrations
of available nitrogen in excess of 100 pounds per acre
in high analysis fertilizers proved to be injurious.
Wilde (1938) recommends available nutrients in the

ratios of 1-2-5. His analysis of the Optimum soil

status for red pine is pH 5.h, base exchange capacity
8.0 m.e. per 100 grams, total nitrogen .12 percent,
available nitrogen 30 pounds per acre, available P205
50 pounds per acre, available K20 150 pounds per acre,
replaceable calcium 1500 pounds per acre, and replace-
able magnesium 300 pounds per acre.

Larson and Stump (1939) experimented with various
levels of nutrients on evergreen seedlings and obtained
better response from.combinations of nitrogen, phos-
phorus, and potassium than from applications of single
elements. In general, it was found that nitrogen
increased t0p growth while phosphorus increased root
development. Auten (19h3) and Lunt (1938) warn against
the use of nitrOgen fertilizers applied at seeding and
recommend a 25 pound per acre application two to four
weeks after emergence.

Roth, Tools, and Hepting (l9h8) found that inorganic
nitrogen fertilizers prevented the severity of little-
leaf disease of shortleaf pine when nitrogen was applied
at the rate of 200 pounds per acre or higher. Shirley
and Meuli (1939) show that the drought resistance of red
pine decreases with the supply of available nitrogen in

the soil, and that the supply of phosphorus tends to
p

increase drought resistance.

Kopitke (l9h1) found that with the level of avail-
able nitrogen adjusted at #0 pounds per acre, the.
effect of potassium on carbohydrate synthesis in red
pine seedlings was most favorable. It was found that
red pine green tissue has the lowest freezing point
(-1.63°C) when available K20 was present in the soil
at 80 pounds per acre. Wilde, Nalbandov, and‘Yu (19A8)
correlate highly fertilized succulent seedlings with a
lower content of alcohol benzine soluble substances
and a higher susceptibility to parasitic organisms.

A relationship between a low specific gravity value

of Jackpine (£1223 banksiana) seedlings and heavy
fertilization in the seedbeds was shown by Wilde and
Voight (1948). The possibility of low specific gravity
as an influence in severity of damage to seedlings by
drought, frost, sunscorch, winter drying, sleet, and
parasites is pointed out.

Mitchell (1939) stresses the importance of a bal-
anced external nutrient supply and correlates this
with internal nutrient concentration and consequent
yields. According to his findings, "forced" seedlings

grown in a well balanced and fertilized environment

are unusually hardy. These seedlings were found to be
significantly better than average stock grown on unfa-
vorable sites. They withstood severe winter freezing
and thawing, and they suffered less early frost injury
than seedlings supplied with a less favorable supply
of nutrients.

Hatch (1935) and‘McComb (l9h3) Point out the value
of mycorrhizal symbiotic fungi to the nutrition of pine
seedlings and the failure of the establishment of coni-

ferous nurseries on prairie soils without these fungi.

HISTORY OF THE SITE

M _S__c_>_i_l Management

The nursery is located west of Hagadorn.Road on
Hillsdale sandy loam.soil. According to Professor
Hudson, Michigan State College farm manager for many
years, the site had not received manure or lhme for
the past ten years. During that period the rotation
was oats, timothy-clover mixture, pasture. Previous
to the last ten years, the rotation was corn, oats,
timothy-clover, pasture. From the standpoint of
nutrients and physical condition, the soil was badly
depleted when the seedlings were started in l9h8.

Previous Treatments 2; thgiNursegy Bed

The first season's growth was normal. During the
first part of June of the second year, when chlorotic
symptoms appeared, the beds were fertilized with ammon-
ium sulfate fed through an overhead sprinkling system
at the rate of 200 pounds per acre. The same treatment
was made again on one half the seed bed early in July.
Neither of the applications had a noticeable effect
on the seedlings. The fertilizer treatments were fol-
lowed by generous sprinklings of water, and the beds
were watered throughout the summer of l9h9 by this

method of overhead watering.

EXPERIMENTAL PROCEDURE
The experimental work discussed in this paper was

started on September 1, l9h9.

Preliminary Investigations

In early September of 19h9, more than 100 borings
were made in the red pine bed to determine whether a
correlation existed between spotty growth and depth
of topsoil, or texture and color of subsoil. No such
correlation was noted. (See Table 1.)

Physical condition of the soil where growth was

 

10

normal was noticeably superior and somewhat sandier.
T0psoil ranged from seven to twelve inches in depth.
Roots of normal seedlings penetrated the subsoil in
most cases investigated*, but the bulk of the root
system was from three to five inches below the surface.

Both surface and subsoil samples were taken on
areas of normal and abnormal growth. The results of
active soil tests (Spurway, l9t9) are shown in Table 2.
Reaction was checked by using a Beckman pH meter with
water-soil ratio Of 2:1. Ten samples each, from repre-
sentative spots in normal and poor growth, were checked.
The results appear in Table 3.

Pigweeds and hardwood seedlings showed signs simi-
lar to those characteristic of manganese deficiency
on certain field crOps, but it is possible that the

chlorosis was due to other causes.

Sgil Preparation

Soil was taken from areas where red pine seedlings
had made poor growth. The areas selected were numerous
enough to represent the entire red pine bed. The soil
was screened through a A mesh screen and thoroughly

mixed. Soil representing normal growth was removed

from exact spots in which normal seedlings occurred

* Based on 10 samples.

11

and was prepared likewise. As a check, Oshtemo loamy
sand from a good red pine site was chosen for one treat-

ment.

‘Pgt Prgparation

Five week old seedlings were used for most of the
treatments. These seedlings were grown for another
experiment from seed planted on August 13, 19h9 in
Plainfield sand surface soil. Their likelihood of
responding to greenhouse conditions during the normal
dormant period seemed greater than that of the 2 year
chlorotic seedlings which had begun to 'harden off"
for the winter. Due to the possibility of reaponse
of the affected chlorotic seedlings, a supplementary
set of pots and treatments, using this material, was
added to the original five week old seedling experi-
ment. Six and seven inch clay pots were used for the
experiment. The five week old seedlings were planted
in six inch pots. Seven inch pots were used for the
affected two year old seedlings. The pots were painted
with asphalt varnish. Each of the two size groups of
pots were brought to equal weight by the addition of
pebbles before the addition of soil. An equal amount

12

of air dry soil was added to each of the pots in each
size group. The six inch pots required 1800 grams of
soil, and the seven inch pots required 2700 grams.

Those pots involving acid peat and those involving

manure treatments were filled by mixing 2/3 of the
required soil by weight with the amount of organic
material necessary to fill each pot. The organic
material (peat and manure) was thoroughly pulverized

and mixed by hand before replacing in the pots. The
fertilizer and sand treatments were thoroughly hand

mixed with soil of their respective pots in like manner.
The mycorrhizae used for inoculation was obtained locally
from a good red pine site, the soil type of which was
Oshtemo loamy sand. Roots of twelve inch red pine trans-
plants were examined for mycorrhizae, and the rootlets
containing these symbiotic fungi were stripped off and

mixed with the soil used for this treatment.

Treatments

Fertilizer treatments were calculated on an area
basis, and the depth of the pots was considered topsoil
depth. The three fertilizer analyses were made from
ammonium sulfate 20-0-0, superphosphate 0-20-0, and

potassium chloride 0-0-50. Minor element treatments

13

consisting of manganese and iron were applied in the
form.of manganous sulfate and ferrous ammonium sulfate.
The fertilizer materials and rates of application are
shown in Table 5. Applications were made at the begin—
ning of the experiment, and a second identical applica-
tion was applied 3 months later. This second fertilizer
application was applied as a nutrient solution immedi-
ately following active soil tests (Spurway). Treatments
(see Table 5) were set up to show the effect of soil
packing (treatment 10), and organic matter amendments

on seedling growth. Acid peat (treatments 11, 12, 13,
23) and stable manure (treatments 1h, 15, 17, 2A) were
pulverized and thoroughly mixed with the soil before

planting.

Description 2; Sgilg

The soil in the affected nursery is classed as
Hillsdale sandy loam. The surface soil is light
brownish to yellowish underlain by yellowish friable
sandy loam.snd gritty clay. The surface may be some-
what stoney. Fertility is classed as medium. The
character of the land is hilly to smooth, rolling,
upland with an original forest type of OakAHickory.

Oshtemo loamy sand was the soil used as a control.

--py.

1b

This soil was selected from an excellent natural red
pine site, and these pots, without treatment, were used
as a check or index with which to compare the various
treatments of the Hillsdale nursery soil. Oshtemo is
described by Veatch (l9hl) as light loamy sands and
sandy loams underlain by pervious sand with small
amounts of clay and gravel. The fertility is regarded
as low to intermediate. This type occurs on level or
pitted dry sandy plains and terraces. The natural
cover was an Open oak-hickory forest with some white

pine in the northern areas.

Planting 222 Culture

All of the pots in the experiment were planted on
September 16, 19h9. The five week old seedlings were
cultured in natural sand prior to transplanting. The
chlorotic two year seedlings were selected in such a
manner that they were a fair representation of the aver-
age size of the affected seedlings and of an average
degree of chlorosis occurring in the red pine bad. They
were taken directly from the nursery bed and transplanted
to the pots.

During the first month of the experiment, outdoor

15

temperatures were unusually high for the season, and the
greenhouse temperature was difficult to control. All of
the seedlings apparently suffered some heat damage.
Many of the two year seedlings died. In view of this
fact, all two year seedlings in the seven inch pots were
replanted on October 12. Partial shade was then provided
during bright days throughout the balance of the early
fall. This was accomplighed by stretching cheesecloth
on a frame above the seedlings. Thus mid-day tempera-
tures were kept at a safe level. Greenhouse tempera-
ture was maintained between 70° and 80° after October 10.
Artificial lighting was used as a daylight supplement for
approximately five hours per day after November 20, until
the end of the experiment on March 27.

Moisture equivalent was determined for all soils,
and soil moisture was maintained as close to this value
as practicable by weight adjustments every week to ten
days. Clay saucers were coated with asphalt varnish
and placed under each pot for watering from below. This
watering from below was alternated with watering from

above.

Measuremepts

At the beginning of the experiment, measurements and

l6

weights of ten average seedlings for each of the follow-
ing classes were taken:

1. 5 week old seedlings

2. 2 year chlorotic seedlings

3. 2 year normal seedlings
The root measurements, top measurements (root collar to
tip), air dry weights, and root-shoot ratios are given
in Table A. These represent initial values at the
beginning of the experiment.

On December 20, 19h9, height measurements above the
soil line were taken of all seedlings to compare treat-
ments by their tOp growth during the first 3 months. The
average length of 10 seedlings by treatment is shown in
Table 5. Since the initial leaves of each of the young
seedlings dried up and were lost in the normal growth
process of these seedlings, some measurements after 3
months appear smaller in the table than the initial
measurements taken at the beginning or the experiment.
Therefore, the new growth which took place during this
period is not satisfactorily revealed by this table at
three mOnths growth. However, a comparison of growth
by treatment is the purpose here, and the differences

in growth are shown by these measurements. While tOp

17

growth was taking place at the terminal bud, thus put-
ting out all new leaves, the original seed leaves dried

up and eventually dropped off.

Harvesting 2; Plants

On March 27, 1950, the plants were carefully removed
in the following manner: The soil was loosened from the
side of each pot with a spatula, and the pot was tapped
gently to allow the contents to drOp from the pot as
one solid ball of soil. The ball was then carefully
broken up to avoid breaking rootlets. The seedlings
were thus taken up with their complete root systems
and placed in beaksrs of distilled water and labeled
according to treatment. By keeping them fresh in this
manner, they were in excellent condition for photo-
graphing and for handling for measurements. After
photographs and measurements were taken, the seedlings
were cut off at the root collar, and the roots and the
tape were placed in separate paper bags according to
treatment. They were dried in a low temperature
(110°?) oven for a week, then allowed to come to
equilibrium at room temperature for another week before

weighing. All weighings were made on an analytical

type balance.

18

Final msasursments in millimeters and weights in
grams were recorded for all treatments. The root and
top lengths as well as the dry tissue weights and the
root-shoot ratios are shown in Table 8a for the young
seedlings. Table 8b shows the same data for the two
year chlorotic seedlings.

A comparison of size of plants as affected by treat-

ment is made in Figures 3 and A.

Soil tests:
In addition to the tOpsoil-subsoil data in Table 1

and the initial soil test and reaction data presented
in Tables 2 and 3 respectively, additional possible
differences in the soils of good and poor growth were
sought.

Additional active soil tests (Spurway, l9h9) for
nitrogen, phosphorus, and potassium were made after 3
months. See Table 6. At the same time, measurements
were taken as stated above. (Table 5). At the end of
the experiment, March 22, active soil tests (Spurway,
19L9) were made for nitrogen, phOSphorus, potassium;

also active and reserve iron and manganese on all treat-

19

ments. See Table 7.

Soil reaction:
Soil reaction was determined with a pH meter.

Results are shown in Table 7.

Moisture equivalent:
Moisture equivalent was determined in duplicate by
the centrifuge method (Vsihmsyer and Hendrickson, 1931).
Determinations were made on the following samples:
1. Hillsdale (poor growth) and 1/3 peat
2. Hillsdale (poor growth) and 1/3 manure
3. Hillsdale (poor growth)
a. Hillsdale (good growth)
5. Oshtemo loamy sand
Samples were placed in moisture equivalent boxes
after screening through a 2 mm. screen. They were then
allowed to become saturated with water for 2h hours and
were permitted to drain for 30 minutes. They were then
subjected to a centrifugal force of 1000 times gravity
by placing in a centrifuge for 30 minutes at 2AAO r.p.m.
The percentage of moisture was then calculated on an

oven dry basis. See Table 83for results.

20

Base exchange capacity and percent base saturation:

Base exchange capacity and percent total base satur-
ation was determined (Schollenberger, 1945) on the poor
growth Hillsdale soil. Results are shown in Table 9.

Organic matter content:

Organic matter content was determined on the Hills-
dale soil samples of good and poor growth for comparison.
The dry combustion method was used (Schollenberger, 19h5)
and organic matter was determined from the organic car-
bon content by use of the factor 1.72h. Values are given

in Table 9.

Phosphorus Determination
Symptoms of phosphorus deficiency (Mitchell, 1939;

Lutz a Chandler, l9h6: Cook & Millar, l9h9) appeared on
a few of the seedlings after approximately 3 to h months
in the greenhouse. Therefore, a total phosphorus deter-
mination was made (Piper, 19A6) based on the weight of
air dry plant material for all treatments of the young
seedling experiment. The determination was made colori-
metrically after dry ashing the plants. Roots and tOps

of each treatment (10 seedlings) were combined and weighed

H.

“a... u..—

21

to the 3rd decimal place on an analytical balance. The
plant material was ashed in a muffle furnace, and the
silica was dehydrated with.HCl over a warm hot plate.
The dehydrated ash was taken up in HCl and diluted to
200 ml. Colorimetric determination was made by use of

a "Lumetron" colorimeter after treating the extract with
ammonium.molybdats and an organic reducing agent. The

results appear in Table 10 as percent total phosphorus.

DISCUSSION AND RESULTS

The benefits derived from the use of soil amendments
and commercial fertilizers in forest nursery management
has long been recognized. (Toumey &'Korstian, 1916;
Wilde, 1916) Nursery soils require particular attention
to avoid depletion of fertility and structural character-
istics. The intensive nature of nursery practice is the
reason for this. Mbst nursery soils are worked to a
very fine state of tilth. This allows the soil to be
more easily eroded. The use of legumes with prOper rota-
tions should be practiced to maintain fertility. Since
trees are taken up by the roots, there are no plant
residues left after a crOp of seedlings is removed frOm a

nursery bed. Therefore, it is reasonable to assume that

’O.

22

nursery management without soil amendments and good
rotaions is soil depleting.

The stunted growth and chlorotic condition of the
2-0 seedlings of the college forest nursery is probably
a result of a combination of adverse soil conditions.

An ideal site for a red pine nursery would demand
loamy sends to sandy loam having excellent drainage. A
soil without radically different composition of its gen-
etic horizons would be in order (Wilde, 1946). Due to
the heterogeneity of the soil profile (see Table 1), it
is possible that variations in drainage, aeration, re-
action, tsxturs, and chemical composition are such as

to cause spotty growth.

Aeration gpd_Drainags

A lack of legumes in past rotations resulted in a
soil of poor aggregation, porosity, and structure. The
very fine sand and clay particles have a tendency to
pack, resulting in seemingly poor aeration and perco-
lation, Due to adverse weather conditions, core samples
for porosity were not obtained. However, spots of good
growth were of noticeably lighter texture. Percolation

rate was checked in the greenhouse by watering 7 inch

23

pots of good and poor growth soil. Percolation was
approximately 20 percent greater in the soil of good
growth. The moisture equivalent values presented in
Table 9 show that organic matter increased water hold-
ing capacity. Values for pest and manure treatments
are rather low, probably due to the raw stats of the
peat and manure used. That good aeration and drain—
age ars important prerequisites to red pine establish-
ment is shown by considering the natural sites on which
this species occurs. Even where competition is elimin-
atsd, red pine will not tolerate poorly drained or poorly
aerated soils.

Due to the poor response of the chlorotic two year
seedling experiment, the remaining results and discus-
sion will refer only to the young seedling treatments

unless otherwise indicated.

ghysicp; Properties

The effect of organic matter treatments to improve
physical prOperties of the soil is reflected by the
excellent growth of the trees where peat was incorpor-
ated with the soil. During the first 4 months of the

experiment, very good growth resulted where manure was

24

used, after which a toxicity devleped which inhibited
growth. Packing the soil depressed growth of the trees.
This is shown by the fact that plants subjected to this
soil treatment produced the second lowest yields of any
in the experiment (Table 10). Total phosphorus was also

extremely low in these plants.

§2_i_l Reaction

The Optimum soil reaction range for red pine has
been determined by Wilde (1946) to be between pH 5.0 and
6.0. This was borne out by the fact that treatment 16,
1/3 alkaline sand (pH 8.0), resulted in the lowest yield
of plant material (Table 8a). These seedlings remained
stunted and unhealthy in appearance throughout the experi-
ment, and they exhibited phosphorus deficiency symptOms
by a purpling of the older needles. The total phosphorus
content of the seedlings of this treatment was rather
low (Table 10). ‘thrs acid peat was used as a physical
treatment (treatment nos. 11, 12, 13), reaction was
lowered by more than one pH unit. Nitrogen added as
ammonium Sulfate solution in treatment 9 contributed
to lowering the pH. (See Table 3 for original pH values;

also Table 7, treatment nos. 19, 3, 4). The relation-'

25

ship between seedling yield and soil reaction is

illustrated graphically by Figure 1.

Effect 2f Organic Matter and Fertilizer

According to Wilde, Optimum organic matter content
should be approximately 2 percent. Results on the
untreated Hillsdale soil (Table 9) correspond favorably
with this figure.

Yield was appreciably higher where fertilizer was
used with peat than in the case of fertilizer used alone
(Tables 8a, 10). Where peat alone was used, yield was
approximately the same as where the soil was untreated
(treatment 19, Table 10). However, the phosphorus
content of the plants was higher, and the root-shoot
ratio and the general appearance of the plants were
more favorable where peat was used (Tables 7, 8a and
10). This was probably caused by the effect of the
peat in lowering soil reaction a full pH unit. Where
fertilizer as 1000 pounds per acre of 10-4-5 was added
with peat, higher yields, higher total phOSphorus, and
improved plant appearance resulted. Still better results"
were obtained where the higher strength fertilizer 10-

12-I2 was applied .

26

Manure treatments resulted in improved growth and
appearance during the first 3 or 4 months after plant-
ing. After February 1, 1950, they began to show signs
of yellowing at the needle tips. This condition became
more acute as time went on. By March 1, some seedlings
thus treated had died. Apparently this condition was
caused by an excess of nitrates resulting in acute tox-
icity. According to Wilde (1946), Optimum available
nitrogen content should be approximately 30 pounds per
acre. According to soil tests (Table 7) made by the
Spurway method of soil testing, nitrates were more than
five times this value. It is also conceivable that in
the process of nitrification, intermediate products
played a role in toxicity. Where peat was used, nitrates A
were high only in treatment no. 12 which had been treated
with 10-4-5, 1000 pounds per acre and where the reaction
was pH 4.5.

Where fertilizer treatments only were used, at 300
pounds per acre the 10-6-6 analysis yielded best results
both for height and weight values. Where the level was
increased to 1000 pounds per acre, average heights and
weights were about equal for 10-6-6 and 10-12-12 analyses.
Values for the 10-4-5 analysis were somewhat lower. Where

only nitrOgen was applied as ammonium sulfate solution,

9‘

P.

27

yield increase was somewhat lower than it was where

the treatment was 300 pounds of complete fertilizer.

Minor Elements
Minor elements (manganese and iron) had little

apparent effect on results (Tables 5, 8a, 10).

Mycorrhizas

Mycorrhizal dsvlepment was best on the roots of
plants grown on the untreated Oshtemo loamy sand (no.
20). Some development of mycorrhizae was also evi-
dent in all pots involving acid peat. The roots of
all plants bearing mycorrhizal fungi were healthy appear-
ing having a light brown color and an abundance of fi-
brous rootlets (treatments 11, 12, 13).. A generally
healthy appearance of these seedlings prevailed through-

out the experiment.

Bass'Exchange
According to Wilde (1946), Optimum base exchange '

capacity for red pine nursery soil is 8.0 milliequi-
valents per 100 grams of soil. Taking this value as a

criteria, 1‘;"would seem that the most desirable method

”-0..-

28

of raising the base exchange capacity would be by the
addition of the proper amount of acid peat. This would
seem especially beneficial since the peat improves soil
physical prOperties and may be used to regulate soil

reaction.

Height Growth
The effect of treatments on the height of the plants

was apparent toward the latter part of the experiment.
Apparently the seasonal dormant characteristics of the
species influenced the time of growth response in spite
of the ideal growing conditions in the greenhouse.

Weights

Mean weights are probably the most satisfactory
measure of growth in young seedlings, since needle drOp
or a rapid rate of needle growth is likely to give tOp
measurements a distorted picture of actual growth gain.
Total yields expressed as air dry weights (Table 10)
and root and tOp weights (Table 8a) were considerably
affected by treatments. The combination of 1/3 peat by
volume and 10-12-12 fertilizer at the rate of 1000
pounds per acre proved far more effective than did any

other treatment.

29

Root-Shoot Ratios

Root-shoot ratios are given in Table 8a with indi-
vidual top and root lengths and weights. Root-shoot
ratio is considered to be an index of planting stock
quality for seedlings two years or older. However,
root-shoot ratios for very young seedlings such as those
used in this experiment would tend to give wide ratios,
since initial growth does not tend toward root develop-
ment as does growth after the first season. Fertilizer
treatments also tend to stimulate tOp growth more than
root growth especially in the earlier months of growth.
This is illustrated by treatments 11, 12, 13, and 20 in
Table 8a. Root-shoot ratio has a tendency to be greater

where nitrogen is limiting (See Tables 7 and 8a).

Egggl’Phosphorus

According to Mitchell (1939), white pine seedlings
showed optimum.growth where their total phosphorus con-
tent was approximately .67 percent. The results of this
experiment show the same to be true of red pine. This
fact is illustrated by the graph in Figure 2 where Opti-
mum growth as measured by dry seedling weight is approx-
imately at the point where total phosphorus is .67

30

percent. It is interesting to note that during the
third and fourth months after planting in the green-
house, treatment numbers 1, 3, h. 9, 10, 14, 16, and
19 showed varying degrees of abnormal purpling of the
needles. This is characteristic of phosphorus defici-
ency symptoms of coniferous seedlings as described by
Mitchell (1939), and Lutz and Chandler (1946).

The extremely high phosphorus content (.98 percent)
of the seedlings grown on untreated Oshtemo sand from an
excellent natural red pine site is worthy of attention.
The excellent deve10pment of ectotrOpic mycorrhizae on
the rootlets of these seedlings may be an explanation to
the high phosphorus content of the plants. According to
mitchell (1934), mycorrhizae tend to deveIOp on the short
roots of seedlings more abundantly in less fertile soils.
It is thought that roots react to nutrient unbalance by
becoming mycorrhizal. .Absorptivs powers of rootlets are
increased manyfold by mycorrhizal development (Mitchell,
l93h; Hatch, 1936; McComb, 1943). Plant analysis shows
nitrogen, phOSphorus, and potassium content of seedlings
having abundant nycorrhizal development to be signifi-
cantly higher than plants having few mycorrhizae. The

difference in total phosphorus in this case was three-

31

fold (Lutz and Chandler, 1946). Mycorrhizal inoculation
has been fOund to be necessary for successful establish-
msnt of coniferous nurseries in prairie soils (Hatch,
1935, 1936). Mycorrhizae were stimulating to growth and
activity of roots, and they allowed roots to absorb more
phosphorus which was apparently limiting to growth in
prairie soils. Mycorrhizal plants contained twice as
much nitrogen and potassium and four times as much phos-
phorus as plants without mycorrhizas (McComb, 1943).
Therefore, it would be reasonable to assume that the high
phosphorus content in the seedlings of treatment 20 (Osh-
temo sand) could be at least partially due to the effici-

ency of mycorrhizal roots in absorbing phOSphorus.

2:9 Chlorotic Seedlings_

A The two year chlorotic seedlings taken from the
nursery bed did not greatly respond to treatment. Little
top growth took place throughout the experiment. Some
root growth took place, however. The effect of nitrOgen
treatments in lowering the root-shoot ratio are evident
in all treatments involving nitrogen application in the
form of manure, peat, and complete fertilizer (see Tables

8b and 4). All root-shoot ratios were narrowed on these

32

seedlings, however, due to root growth only.

SUMMARY AND CONCLUSIONS

A greenhouse eXperiment was set up to determine the
cause of chlorosis and the stunted growth which occurred
in the red pine forest nursery of Michigan State College.
Soil from the affected area was prepared and placed in
clay pots. Five week old seedlings were used for one
part of the experiment, and two year old chlorotic seed-
lings from the affected nursery were used for the other
part.

Treatments consisted of various levels of complete
fertilizer, peat, manure, sand, and combinations of peat
and fertilizer. Little response occurred from.the two
year old chlorotic seedlings. The "new" five week old
seedlings responded very well to treatment. In the case
of fertilizer treatments only, the 10-6-6 analysis gave
best results in both height growth and in yield. Acid
peat with 10-12-12 fertilizer at the rate of 1000 pounds
per acre gave the highest yield. Plants appeared vigor-
one and healthy where peat was used. Peat applied at the '
rate of 1/3 by volume lowered the reaction by more than

a full pH unit. A correlation between pH and yield was

33

noted. A pH range of 5.5 to 6.5 appeared to give best
results. Manure treatments deve10ped toxic symptoms
after the fourth month of the experiment. This was
probably due to the effect of excess nitrates and inter-
mediate compounds produced in the process of nitrifica-
tion. Depressing effects on yield by high reaction was
reflected by the treatments involving alkaline sand.
Abundance of available nitrogen decreased root-shoot
ratios due to its effect in stimulating t0p growth.

Oshtemo loamy sand was used as a control, and it
produced the second highest yield. Mycorrhizal devel-
opment was excellent on seedlings grown in this soil,
and some mycorrhiZae were present on all treatments in-
volving peat.

Phosphorus deficiency symptoms developed on several
of the treatments which received low fertilizer applica-
tions. No apparent deficiency was evident where peat was
used. A good correlation between total phosphorus con-
tent of seedlings and yield was noted. High phosphorus
content of seedlings grown on Oshtemo loamy sand may be
correlated with exceptional mycorrhizal deve10pment and
the ability of these symbiotic fungi to assist in taking

up phosphorus from the soil.

The possibility of phosphorus deficiency as well as

31+

the need of better physical prOperties and a somewhat
lower reaction for the problem soil was presented by

the results of this experiment.

Table 1.

Number of
boring

1*
2**
3*
1+**
5*
6**
7*
8**
9*
10**
11*
12*!!!
13*
lh**
15*
16**
17*.
18**

depth
6"
7»
7n
6n
8"
11"
ll"
10"

10"

9.
10"
10"
11"
10¢
11"
10"
11"

35

TOpsoil

* Areas of poor growth

** Areas of good growth

texture

sand
sand
sand
sand
sand

sand

loamy sand

clay

clay loam

clay

loamy sand

sand

loamy sand

sand
clay
sand
clay
clay

dubsoil

A comparison of soil in areas of good
and poor growth.

(Soil color-texture-growth relationship)

color

light

gray
light

gray

light
light

light

medium

light
light

brown
brown
brown
brown
brown
br own
brown
brown
brown
brown
brown
brown
brown

brown

medium brown

medium gray

mottled gray

yellow brown

Table 1.

Number of
boring

19*
20**
21*
22**
23*
2h**
25*
26**
27*
28**
29*
3020“
31*
32**
33*
31P**
35*
36**

* Areas of poor growth

** Areas of good growth

continued

TOpsoil
depth

8»
9n
10"
ll"
10"
ll"
8"
12"
7w
1h"

ll"
7"
ll"
8"
ll"
8"
10"
9w
1 O"
10"

texture
sand
sand
loamy sand
sand
loam
sand
loamy sand
sand
loamy sand
clay
sand
sand
sand
sand
sand
loamy sand
loam
sand
sandy loam
loam

sandy loam

subsoil

color
med ium br own
gray
light brown
medium gray
medium brown
gray

medium brown
gray brown
light brown
gray mottled
light brown
medium gray
gray

medium gray
light brown
gray brown
light brown
yellowish
medium brown
medium gray

medium brown

37

Table 1. continued

Number of Topsoil
boring texture

AO** 9" sand
hl* l2" sand
h2** ll" sand
h3* 9" sand
hh** 9" loam
45* ll" sand
L6”* 9" sandy loam
47* ll" sand
h8** 9" sand
h9* 8" sand
50** 7" sand
51* 10" sand
52** ll" loamy sand
53* ll" sand
5L** 8" sand
55* 10" sand
56** 8" sand
57* 9" sand
58** ll" sand
59* 9" sand
60** 9" sand

* Areas of poor growth
** Areas of good growth

subsoil

color
light brown
medium gray
gray
light gray
light gray
light gray
gray brown
light gray
yellow brown
reddish brown
gray brown
yellow brown
gray

medium gray
gray brown
medium gray
gray
red brown
medium gray
reddish—yellow

light gray

Table 2.

Nitrates
Iron
Phosphorus
Potassium
Calcium
Magnesium
Manganese
Chlorides
Sulphates
Nitrites
Aluminwm

38

Normal Growth

Surface

Ppm.

h
0

trace

3
so
6
0
3o
20
0
0

Subsoil
Ppm.

3
O

3/h

100

30
20

Results* of active soil test (Spurway Method)
of soil from nursery bed at the beginning
of the experiment, September 16, l9h9.

Poor Growth

Surface
Ppm.

10
O

trace

no

30
20

Subsoil
Ppm.

5

* All values are in parts per million in soil extract-

soil-water ratio lzh.

39

Table 3. Beckman pH meter readings on soils from
areas of good and poor growth.

Sample

No. Normal Growth Poor Growth
1 6.1 6.7
2 6.2 6.1
3 6.7 6.h
h 6.8 6.6
5 6.9 6.0
6 6.5 6.h
7 6.h 6.0
8 6.3 6.0
9 6.5 6.5

10 6.9 6.2

LO

Table A. Mean lengths* and weights** and root-shoot
ratios of seedlings at the beginning of the
experiment, September 16, l9h9.

Age of

Seedlings Length (mm.)
Root TOp

New*** 39.7 60.9

2 yr. stunted 226.5 92.5
2 yr. normal 2L2.1 163.5

* Mean value-~10 items
** Total value-~10 items
*** 5 weeks old

Weight (g.)

Root TOp

.02 .12
1.2h 2.72
3.h5 10.n2

Root-shoot
Ratio

.18
.L5
.33

hl

Table 5. Heights* of seedlings after approximately
3 months in the greenhouse, December 20,

New Seedlings (6" pots)

No. Treatment Height (mm)
1 lO-h-S, 300 1bs./A u1.u
2 10-6-6, 300 lbs./A a9.o
3 10-12-12, 300 lbs./A 4a.9
a 10-u-5, 1000 1bs./A 40.0
5 10—6-6, 1000 1bs./A ha.3
6 10-12-12, 1000 1bs./A L3.5
7 lO-L-S, 1000 lbs./A and Mn, 100 lbs./A u3.u
8 10-a-5, 1000 1bs./A and Fe, 100 lbs./A h5.0
9 (NHA)ZSOL solution, 200 lbs./A h3.1

10 Soil** firmly packed ' no.3

11 1/3 peat 39.9

12 1/3 peat and 10-h-5, 1000 1bS./A h1.0

13 1/3 peat and 10—12-12, 1000 1bs./A b7.h

it 1/3 manure . no.6

15 1/3 manure and 0-20-0, 250 lbs./A and

0-0-50, 100 lbs./A h3.6

16 1/3 sand (pH 8.0) 34.8

17 l/h sand (pH 8.0) and l/h manure 3h.6

18 Soil of good growth, no treatment h5.8

* Mean value-~10 items
** "Soil” refers to soil of poor growth unless other-
wise indicated.

A?

Table 5. continued

No. Treatment Height (mm.)
19 Soil of poor growth, no treatment hl.h
2O Oshtemo loamy sand, no treatment h5.3

Chlorotic 2-O* Seedlings (7" pots)

21 lO-h-S, 1000 lbs./A 92.8
22 1/3 sand (pH 8.0) 82.6
23 1/3 peat . 81.3
2h l/h manure ,77.1
25 10-12-12, 1000 1bs./A 75.3
26 1/3 Oshtemo sand and mycorrhizae 72.2
27 10-5-5, 1000 1bs./A and Mn, 100 lbs./A 6a.u
28 10-4-5, 1000 lbs./A and Mn, 100 1bs./A

and Fe, 100 lbs./A 78.7
29 Soil of poor growth, no treatment 68.5
30 Soil of good growth, no treatment 190.3

* 2 years in seed bed; 0 years in transplant bed

L3

Table 6. Active soil tests (Spurway) after 3
months in the greenhouse

Treatment Parts per million in soil extract*
N P K iMn Fe
io-t-S, 300 1bs./A 0 0 t 0 0
10-4-5, 1000 lbs./A trace trace is 0 0
10-12-12, 1000 lbs./A 0 trace 7 o o
io-t-s, 1000 1bs./A
and Mn 100 lbs./A trace trace 6 O O
(NHL)ZSOQ solution,
200 lbs./A 0 trace t~ 0 0
Soil packed O 1/2 A O 0
1/3 peat 2 O O O
1/3 peat & 10-12-12,
1000 1bs./A 2 trace 20 0 0
1/3 manure lO l/2 25 O 0
good growth soil-none 15 O 5 O 0
poor growth soil-none O O 6 O O
Oshtemo (natural site)
none 0 O 3 O O

* Soil-water ratio lzh

Table 7.

Treat.

NO.

\OG)\}O\\J'I(>\ANI~'

HI—‘i—‘l—‘I—‘I—‘HH
\jome'wNI—JO

hh

Results of soil tests (Spurway) taken at the

end of the experiment, March 22, 1950.

Ppm. in soil extract--soil water ratio l:h

00002

10
25

GOOD

tr.

25
25

Active

W’U

h‘ are uu

25
E

H
00‘

\iocwvuwrmmm

k' (0 N ha
04>me

OOOOOOOOOO

tr.

tr.

tr.

tr.

000000

('9'
H
.

O O O 0 I39“ 0 O O O O

by
0

OOOOOOOOOOOOOOOOO

Reserve
Mn Fe
1 2
1 2
l 2
1 2
1 2
l 2
2 2
l 2
1 2
l 2
tr. 1
tr. 5
l h
tr. tr.
tr. tr.
tr. tr.
tr. tr.

DH

6.50
6.02
6.50
5.32
5.35
5.30
5.58
5.50
5.59
7.05
6.15
b.55
5.75
7.05
6.75
8.02
7.21

h5

Table 7. continued

Active Reserve

Treat.

No. N P K NH3 Mp Fe Mn Fe pH
18 0 tr. 2 o 0 0 tr. tr. 6.50
19 0 i 3 o 0 0 tr. tr. 7.10
20 o a 2 o o 0 0 t 6.15
21 5 1 3 O O 0 tr. tr. 5.22
22 0 tr. 1 0 0 0 tr. tr. 8.00
23 0 2 2 2 0 0 tr. 1 6.35
24 15 as 15 2 0 0 tr. tr. 7.15
25 2 a 4 tr. 0 0 1 2 5.38
26 0 tr. 5 tr. 0 0 tr. tr. 6.80

(’1‘

(7‘ :10 Fightz r [r1 .

. . He‘ll.

L6

Table 8a. Root and tOp measurements*, weights**, and
root-shoot ratios of new seedlings at the
end of the experiment, March 28, 1950

 

 

 

Length (mm.) Weight (g.) Root-shoot
Ratio
Treat. TOp Root Top Root
NO.
1 59 186 .h26 .531 1.25
2 7h 221 .652 .662 1.03
3 62 152 .508 .7h8 1.h7
h 62 192 .566 .568 1.00
5 77 200 .803 .731 .91
6 80 187 .935 .810 .87
7 62 181 .622 .688 1.10
8 62 205 .590 .790 1.3h
9 58 201 .506 .722 1.h2
10 5h 13h .hll .h98 1.21
11 65 229 .516 .532 1.03
12 93 183 .886 .h90 .55
13 113 177 1.331 .68A .51
1h 55 156 .h27 .355 .83
15 66 15h .598 .519 .87
16 5h 229 .h15 .h80 1.15
17 56 197 .t55 .h8l 1.03
18 70 251 .650 .798 1.22

* Mean value~~10 items
** Total value-~10 items

Table 8a.

Treat.
No.

19
20

#7

continued

Length (mm.) Weight (g.)
Top Root Top Root
56 203 .h60 .627
98 279 1.028 1.050

Root-shoot
Ratio

1.36
1.02

Table 8b.

Treat.
No.

21
22
23
2h
25
26
27
28
29
30

Root and t0p measurements*, weights**,
root-shoot ratios of 2 year affected
seedlings taken at the end of the experi-

A8

ment, March 28, 1950.

Length (mm.)

TOp

98
87
89
87
92
85
89
85
93
162

306
223
309
2h6
297
307
27h
2A9
331
#91

*-¥ean value--10 items

Root

** Total value--lO items

Weight (g.)

Top

b.537
3.590
3.958
3.162
3.328
2.503
3.351
3.650
2.820

10.730

Root

3.772
3.63h
3.372
2.525
2.578
2.952
2.30h
2.251
2.750
9.3h2

and

Root-shoot

Ratio

.83
1.01 .
.85
.80
.77
1.18
.68
.62
.97
.87

L9

Table 9. Moisture equivalents of greenhouse soil
treatments; Organic matter, base exchange
capacity and percent base saturation of
nursery soil used in experiment.

Soil M.E. 73 0.11. B.E.C. 95 Base
(m.e./100 g.) Saturation

Hillsdale
(poor growth) 11.6 2.2 7.03 85
Hillsdale
(good growth) 15.8 2.9 -- --
1/3 peat 15.0 -- -- --
1/3 manure 13.9 -- -- __
Oshtemo

loamy sand 5.8 -- —- --

50

Table 10. Overall lengths*, total weights**, and total
phosphorus content of new seedlings at the
end of the experiment, March 28, 1950.

Treatment overall Total % total
length weight phosphorus
(mm.) (8.)
10-1-5, 300 1bs./A 245 0.957 .08h
10-6-6, 300 1bs./A 295 1.31t .263
10-12-12, 300 1bs./A 21L 1.256 .128
10-h-5, 1000 lbs./A 25h 1.13t .136
10-6-6, 1000 1bs./A 277 1.737 .3ho
10-12-12, 1000 1bs./A 267 1.7t5 .532
lO-h-S. 1000 lbs./A and
Mn, 100 1bs./A 2&3 1.310 .220
lO-L-S, 1000 lbs./A and
Fe, 100 1bs./A 267 1.389 .260
(NH ) 30 solution
a 2 h 200 1bef/A 259 1.288 .170
Packed soil 188 0.909 .085
1/3 peat 7 29h 1.0h8 .189
1/3 peat and 10-1-5,
1000 1bs./A 276 1.376 .216
1/3 peat and 10-12-12,
1000 lbs./A 290 2.015 .665
1/3 manure 211 . 0.782 .137
1/3 manure & 0-20-20,
250 1bs./A; 0-0-50,
100 1bs./A 220 1.117 .282

* Average value--1O items
** Total value-—10 items

51

Table 10. continued

Treatment Overall
length
(mm.)

1/3 sand (pH 8.0) ‘ 283

k sand (pH 8.0) and

A manure 253

Good growth soil--
no treatment 321

Poor growth soil--
no treatment 259

Oshtemo loamy sand--
no treatment 377

Total
‘weight
(8.)

0.895

0.936

l.hh8

1.087

2.078

$ total
phosphorus

.lh3
- .180
.h05
.152

.980

0L

f HUI Also

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3.9!

 

53

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Figure 3. Seedlings at the end of the experiment
showing the effect or the treatments on

root and top growth.

55

  

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Figure A. Seedlings at the end of the experiment
showing the effect of the treatments on

root and tOp growth.

56

REFERENCES

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