CHEMICAL AND BIOLOGICAL EFFECTS OF LIME APPLICATION
TO BOG LAKES IN NORTHERN MICHIGAN

by
Thomas Frank Waters

A THESIS
Submitted to the School of Graduate Studies of Michigan
State University of Agriculture and Applied Science
in partial fulfillment of the requirements
for the degree of
DOCTOR OF PHILOSOPHY
Department of Fisheries and Wildlife
1956

P roQ uest Num ber: 10008677

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TABLE OF CONTENTS

INTRODUCTION.......................................... ...
Description of area
• • • • • • ♦ # * • 5
METHODS
.............................................
5
Description of lakes
Starvation Lake,
• • • • ................... 6
Timijon Lake#
# • # # • • • • • • • 8
Juanita Lake.
• # . . . # • • . . . 8
Irwin L a k e ...................
10
Grant1s L a k e ..................................... 10
Preliminary laboratory experiments
Application of lime

#

•

•

Method of sampling.

#

•

•

Chemical sampling *

•

•

•

♦

.

•
•

.

•
.

•

*

.

.

•
•

•
.

14
#
•

#

•
•

.

•

1
.
1

Biological sampling
• •
Net phytoplankton * • • • • ♦ • # . * 1
Z o o p l a n k t o n ................• • • • • • •

4
16
7
19
9
20

RESULTS AND DISCUSSION • .
20
Preliminary laboratory e x p e r i m e n t s .............. 20
Release of phosphorus.
• # • • • • • • 2 0
Calcium adsorption#
• • • • # . # • . 2 1
Field experiments . • . . # . • • • • • 2 7
Starvation Lake*
• • • • • • • • • • 2 8
Effects in epilimnion.
* • • • • • • 2 8
Effects in hypoliranion • • • • • • • 3 1
Effects upon phosphorus concentration
• *33
Calcium adsorption by mud
• • • • • •
34
Biological effects#
• • • • • . • • 3 6
Timijon Lake# • • • • • • • * • . . 3 9
Effects in epilimnion*
.............. 39
Effects in hypoliranion • • • • • • • 4 5
Effects upon phosphorus concentration
• • 47
Calcium adsorption by mud
♦ • • • • •
48
Biological effects*
. • • • • • • • 4 8
Juanita Lake.
• • ♦ • • • • • • • • 4 9
• • • • • • • • 4 9
Chemical conditions
• • * • • • • 5 5
Biological conditions.
Irwin Lake
• • • • • • • • • • * • 5 8
Chemical conditions
• • • • • • • ♦ 5 8
Biological conditions*
• • • • • • * 6 0
Grant’s Lake.
* • • • • • • • • ♦ • 6 6
Chemical conditions
• • • • • • • • 6 6
Biological conditions*
• • • • • • • 6 6

GENERAL DISCUSSIONS AND C O N C L U S I O N S ......................... 71
Effects of alkalinity upon phytoplankton production*
• 71
Release of phosphorus from lake soils
Color relations in bog waters *

•

•

•
•

•
•

•
•

•
•

•
•

Comparison of colored and clear-water bog lakes •
Chemical conditions
• * • • • • • • • •
Biological conditions.
• • •

•
•

77
8

2

• • 86
• 8 6
87

S U M M A R Y ...................................................... 88
A C K N O W L E D G E M E N T S .............................................91
A P P E N D I X ...................................................... 92
LITERATURE C I T E D ........................................... 110

INTRODUCTION
In the past, considerable attention has been paid by workers
in many fields of research to bogs and bog lakes*

The unique set

of environmental conditions represented by a bog lake, with its comp­
arative isolation from outside influences, is particularly attractive
to ecologists in several biological fields.

From the limnologist1s

viewpoint, the bog lake, with its acid and extremely soft-water con­
ditions and its encroaching bog mat, presents an object of study which
is totally different in many respects from all other bodies of water,
and workers in this field have studied bog lakes in many investiga­
tional researches (Welch, 1936a, b, 1938a, b, 1945, Jewell and Brown,
1929, Gorham, 1931, and others).

However, despite the abundance of

these lakes in certain regions of northern Michigan and Wisconsin
where sports fisheries are extremely important, it has not been until
recently that this type of lake has been included in developmental
research programs aimed at proper management of bog lakes as sport
fishery resources*
Several reasons are apparent*

Generally, interest shown in

utilizing bog lakes as fishery resources has been low*

Bog lakes are

often very small and perhaps unnoticed; many do not contain desirable
fishes (or perhaps none at all); and they cannot be waded nor approached
easily either by foot or boat.

But one of the greatest deterrents to

giving the bog lake an important position among our fishery resources
is its extremely low biological productivity.
It has been often observed that a correlation exists between the
biological productivity of lakes and the alkalinity of their waters.
Ball (1945), reporting on the results of poisoning lakes in Michigan,
divided the lakes which were poisoned and for which complete fish re­
coveries were attempted into three categories of methyl-orange alkali-

nity and observed that the more alkaline categories contained higher
average standing crops of fish.

It has been suggested, furthermore,

that the alkalinity of lake waters be used as an index to productivity
Moyle (1949), in a study of this and several other suggested indices
of lake productivity in Minnesota, compared total alkalinity and pro­
ductivity (based on pond fish yield and test-net catches in lakes) and
reported that 40 ppm. total alkalinity appeared to be a valid separa­
tion point between soft-water (less productive) and hard-water (more
productive) lakes.

He concluded that total alkalinity, along with

total phosphorus, appeared to be the most valuable of the indices
studied.

Since the existence of causal relationships is a possibility,

a valid phase of developmental research appears to be an investigation
of the use of alkalinizing compounds,

such as lime, to increase the

alkalinity of lake waters and thus, possibly, biological productivity.
It is considered appropriate here to define the terms "alkalization" and "allcalinization, " as used in this report.

"Alkalization"

is used as being synonymous with "lime application," and is not meant
to imply that the water is necessarily made more alkaline, even though
such an effect may be expected; the term "alkalinization" is reserved
for the purpose of denoting an actual increase in alkalinity.
It was the purpose of this research program to test the use of
lime to increase biological productivity in bog lakes, and to report
upon the chemical and biological effects of such treatment.
program permits the attempt to answer several questions:

Such a

(1) What

causal factors are involved in the higher production observed in more
alkaline waters?

(2) What direct effects, if any, would there be upon

organisms living in the lake waters?

(3) What would be the effect of

changing pH levels upon the availability of nutrients?

(4) What fac-

3
tors would reduce the alkalinizing effects of lime treatment, such as
calcium loss due to ion-exchange at the mud-water interface?
In agriculture, the use of lime on acid soils is widely recognized
as a very necessary management practice.

The above questions, posed

about an aquatic medium, have been amply answered about terrestrial
soils*

More alkaline conditions in soil favor the activities of de­

composition bacteria, thus mineralizing nutrients from unavailable
organic forms.

Phosphorus availability depends upon the level of soil

reaction, undesirable levels being at both ends of the pH scale*

Di­

rect effects of soil reaction upon soil organisms are known, particu­
larly disease-causing organisms.
However, the use of lime in aquatic production in this country
is very small compared to its terrestrial use*

In Europe, where the

pond production of carp has been practiced for centuries, lime has been
used to produce increased fish yields (Neess, 1949)*

Lime has been

added with other fertilizers in pond and lake fertilization experi­
ments, both as a fertilizer itself (Juday and Schloemer, 1938), and
for offsetting acidifying effects of ammonium fertilizers (Swingle and
Smith, 1939)*
In Michigan, a small application of hydrated lime was made to a
soft-water, non-bog lake in 1943, expressly for the purpose of in­
creasing alkalinity, and was again treated in 1945 (Ball, 1947), this
time with limestone, neither treatment effecting a change in alkalinity;
the same lake was again incorporated in an alkalization program of re­
search in the years 1952 through 1955 (Ball and Waters, MS), where
changes in alkalinity resulted and increases in standing crop of net
phytoplankton were observed*
The use of lime in treating bog lakes was investigated in Wisconsin

4
in 1948 by Hasler, Brynildson and Helm (1951).

They offered several

suggestions as to the mechanisms by which productivity may be increased
by liming in colored bog lakes.

The c o l o r -caused by humic colloids —

would be flocculated and precipitated through combination with calcium,
thus permitting deeper light penetration and increasing the volume of
the trophogenic zone.

Secondly, alkalinization of the water would

mobilize nutrients which are fixed in unavailable organic forms in
the lake soils.

Finally, a higher concentration of bicarbonates would

result, offering available carbon dioxide to plants for photosynthesis
in higher concentration.

Following their experiment, they reported

that the lime treatment resulted in clearing the color of the water
and improving oxygen conditions for trout in the deeper, cooler levels
of the lakes, but showed no striking differences in nutrient content
of the water*

It was later reported (Johnson and Hasler, 1954) that

no apparent increase in production or carrying capacity resulted.
In the summer of 1953, the present research program was initiated
upon several bog lakes in the Hiawatha National Forest in the upper
peninsula of Michigan.

The program consisted essentially of gathering

pre-treatment data on limnological conditions, applying hydrated lime
to the lakes, and obtaining data on post-treatment conditions to evalu­
ate the changes effected.
Three postulates were made regarding the mechanisms by which
biological productivity might be increased by the use of lime, and
the sampling program was designed to test these postulates:

(1) a

greater concentration of bicarbonate alkalinity, offering more avail­
able carbon dioxide for photosynthesis, would result; (2) phosphorus
would be released either by increased decomposition activities or
through ion-exchange phenomena in the mud; and (3) the colloidal or-

5
ganic color would be decreased by flocculation and precipitation caused
by combination with calcium.
Description of area
The region of the Hiawatha National Forest in Michigan's upper
peninsula was selected for this series of experiments due to its
abundance of small bog lakes.

This region lies near the western edge

of the Lake Superior Lowlands, of the Central Lowland province, and
has a relatively deep layer of £ acial drift underlain with high-calcium
sedimentary rock (Lobeck, 1950).

The sedimentary rock immediately below

the glacial drift in the Hiawatha National Forest is limestone (Bergquist, 1937), and it would be expected that the glacial drift contain
a large proportion of limestone and limestone flour.

This is evidenced

by the alkaline characteristics of deep wells, springs and spring-fed
lakes in the area, and by the presence of limestone in an area gravel
pit; seepage lakes, however, including bog lakes, are acid in nature
and very soft.

The region contains several large outwash plains and

some morainic areas from the Newberry and Munising moraine systems*
The outwash plains, particularly, are dotted with many small glacial
lakes, known as pit lakes; several appear in morainic regions where
they are known as kettle lakes; still others are shared by the edges
of both outwash plains and moraines and are then known as fosse lakes.
METHODS
Description of lakes
A limited survey of small bog lakes in the Hiawatha National
Forest was made during the summer of 1953 and five of these lakes
were selected for experimental purposes.

Among the five lakes were

three colored bog lakes and two clear-water bog lakes.

They were se­

lected for certain desired characteristics, including similarity of

size* chemical and biological conditions, and accessibility.
characteristics were constant for all lakes selected.

Several

All lakes were

completely surrounded by an encroaching, acid, sphagnum-leatherleaf
bog mat.

In addition to Sphagnum and leatherleaf (chamaedaphne caly-

culata, Muench), many other typical bog plants comprised the mats of
all lakes, including several ericaceous shrubs such as the cranberry,
Vaccinium macrocarpon Ait.; bog rosemary, Andromeda glaucophylla Link;
bog laurel, Kalmia polifolia Wang,; and labrador-tea, Ledum groenlandicum Oeder; several sedges, including cotton-grass, Eriophorum spp,;
Carex spp,; and Rhynohospora alba (L,) V a h l ; also the pitcher plant,
Sarracenia purpurea L.; sundew, Drosera rotundifolia Wang,; and an
orchid, swamp-pink, Calopogon pulchellus (Salisb.) R. Br.

Aquatic

vegetation was scant in all lakes and consisted of Nuphar and/or
Utricularia,
All lakes had approximately the same bottom types, consisting of
brown fibrous peat near the mat and a fine, gelatinous pulpy peat of
brown color in the deeper areas of the lakes; no exposed

sand or

other inorganic bottom was found in any of the lakes.
Starvation Lake
This was a colored kettle lake In the Newberry moraine system.
Figure 1 shows a map of marginal outline and bottom contours.
slopes were present on the east and west sides.
forested, even out upon the bog mat.

Steep

All sides were densely

Spruce and tamarack were found

on the mat, with some white pino along the landward edges of the mat.
Poplar, cherry and pine were found on the slopes.

The encroaching

mat was of moderate width on all sides, while a large, connecting bog
mat area lay near the south end, apparently draining into the lake,
A large, active beaver lodge was present on the edge of the bog mat.

7

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Starvation Lake was 1*7 acres in size with a volume of 32*6 acre-feet
and a maximum depth of 42 feet*
nets and "by angling,

Despite repeated attempts with gill

no fish of any kind were ever taken, and none

were ever observed nor reported by others*

See Plate I.

Timijon Lake
This was a colored pit lake in an outwash plain, possessing only
shallow slopes on all sides*

It was separated from a moderately sized

stream, the Big Indian River, by only a narrow dike of glacial drift
of about 50 feet wide; however, no connection between the lake and
stream was apparent nor was one believed to exist*

The lake was

lightly forested on all sides by poplar and birch, and a very few
white pines*

Spruce and tamaracks were found on the bog mat*

The

mat was of moderate width on three sides, while an extensive mat area
was found on the southeast side*

Timijon Lake was 2.0 acres in size

with a volume of 43*1 acre-feet and a maximum depth of 42 feet.

Yel­

low perch, Perea flavescens (Mitchell), were found to be very common;
northern brown bullheads, Ameiurus nebulosus nebulosus (LeSueur),
were also found.

See map in Figure 2, and plate II.
Juanita Lake

Juanita Lake was a colored fosse lake with parts of the Newberry
moraine on the north and west sides and outwash plain on the remainder
of the shoreline*

Slopes on the moraine side were heavily forested

with white nine, birch and poplar.

An extensive bog mat area was found

on the southeast and north sides; the north mat included a small, very
shallow pond, separated from the lake by the mat.

Spruce and tamarack

were found on the bog mat, while Utricularia was very common along the
edges of the bog mat.

Juanita Lake was 1.4 acres in size with a volume

of 35.1 acre-feet and a maximum depth of 34 feet.

Yellow perch, Perea

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Map of Timijon Lake

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10
flavescens (Mitchell), were found to be common; bullheads were reported
in the lake but the species was unknown.

See Figure 3 and plate III#

Irwin Lake
This was a clear—water fosse lake, having the Newberry moraine
on the east side and outwash plain on the remainder#

It was densely

forested on all sides except the west; white pine and birch were pre­
dominant.
tamaracks#

The bog mat was narrow on all sides and held few spruce and
Aquatic vegetation consisted of a few patches of Nuphar#

Irwin Lake was 10 acres in size and had a maximum depth of 38 feet#
The lake originally contained yellow perch, Perea flavescens (Mitchell),
but had been poisoned and restocked with brook trout, Salvelinus fontinalis fontinalis (Mitchell)*

See map in Figure 4, and plate IV#

Grant?s Lake
This was a clear-water bog lake of the fosse type, having parts
of the Newberry moraine on the north and west sides, and outwash plain
on the remainder#

The surrounding land was moderately forested on all

sides with poplar and birch primarily and a few white pines*

Only a

few spruce and tamaracks were found on the bog mat, and the aquatic
vegetation consisted of Nuphar, which was relatively common#
mat was narrow on all sides#

The bog

This lake was h rger and shallower than

any of the other lakes and did not stratify thermally except in a small
depression of deeper water#

This lake, originally containing yellow

perch, perca flavescens (Mitchell), had been poisoned and restocked
with rainbow trout, Salmo gairdnerii irideus Gibbons, and brook trout,
Salvelinus fontinalis fontinalis (Mitchell).

Grant*s Lake was 13.8

acres in size and had a maximum depth of 30 feet#
and Plate V •

See map in Figure 5,

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14
Preliminary laboratory experiments
Two postulates were investigated in the laboratory before any
applications of liming materials were made to the lakes.

The first

was that phosphorus would be released from the bottom organic deposits
upon the application of lime; the second was that the calcium from the
applied lime would enter into ion—exchange reactions at the mud—water
interface and be lost from the lake water.

These postulates were

tested by conducting experiments involving the addition of hydrated
lime at known rates to bottles containing mud and water which had been
collected from the three colored bog lakes, and testing the water for
dissolved phosphorus and dissolved calcium.

A separate series of ex­

periments was conducted for the testing of each postulate.

Detailed

procedures are included in a later section with the results.
Application of lime
For the application of the lime to the lake waters, a raft-andpump system was employed which mixed the lime with water and discharged
the mixture, which was a white, milky suspension, at the surface of
the water.

The apparatus consisted essentially of a flat raft ten

feet long and six feet wide, supported in back by the edge of the bog
mat and in front by two floating 50-gallon drums.

Through a rectangular

hole in the raft was lowered a plywood mixing tank which allowed the in­
take of water at the bottom of the front side of the tank; a hopper
was constructed on top of the mixing tank through which the lime was
introduced.

The intake hose of the pump was placed in the mixing tank,

where both lime and water were taken up, carried through the mechanism
of the pump and through the exhaust hose of the pump, to be finally
discharged into the lake (see Plate VI).
Lime application was made to two of the colored bog lakes, re-

a

15
taining the third colored lake for a control.

The two clear-water

bog lakes were retained for comparative studies, particularly should
the lime have resulted in clearing the color from the treated lakes.
Each of the two treated colored lakes received two separate lime
applications, one during the early summer while thermal stratification
was present and the second during the autumn after stratification had
been destroyed.

The summer application to Starvation Lake commenced

on July 10, 1954, and ended July 13, 1954; a total of 1650 pounds was
applied, which amounted to a rate of 50 pounds per acre-foot, on the
basis of the entire volume of the lake (excluding the bog mat).

Liming

commenced on Timij oh Lake on July 19, 1954, and continued throughout
the remainder of the summer to August 28, 1954, on which date the last
application was made.

A total of 1600 pounds was applied for a rate

of 37 pounds per acre-foot, on the basis of the entire volume of the
lake (excluding the bog mat).
The difference between the methods of applying lime to Starvation
and Timijon lakes during the summer was essentially as follows:

Ah

application of 50 pounds per acre-foot was made as rapidly as possible
in Starvation Lake, producing large and precipitous changes in chemical
conditions, particularly the elevation of pH*

The application of lime

to Timijon Lake was made without reference to the rate applied, but
rather to the change in pH produced; it was made at such a rate as to
keep the pH of the surface waters near neutrality.

In both lakes,

during the summer application, discharge of the lime-water suspension
was made near shore immediately in front of the raft (see Plate V I ).
The autumn application to Timijon Lake was made on October 17,
1954, when the lake was nearly homothermous, and consisted of 2700
pounds.

The autumn application to Starvation Lake was made on October

16
19, 1954, consisting of 1650 pounds.

These applications brought the

total rates of application up to 100 pounds per acre-foot for each
lake.

For the autumn applications, the lime-water discharge was made

near the center of the lake with the use of hoses suspended by buoys
(see Plate VII).
The lime used was high—calcium hydrated lime, furnished by the
Cutler—Magner Company, Duluth, Minnesota,

An analysis of the material

showed a determination of calcium of 95% (as Ca(0H)2 <)

neutralizing

power of 90% Ca(0H)2 (122% CaCO^).
Method of sampling
A periodical sampling program was initiated on all lakes immedi­
ately after selection and mapping of the lakes during the summer of
1953,

The three colored bog lakes— Starvation and Timijon, which were

treated, and Juanita, the control— were sampled every ten days.
two clear-water

bog lakes were sampled three times

The sampling station was selected

so as to be

est part of the

lake, usually near the center.

At

lake a buoy was

anchored securely, and at the time

The

per year.
at or nearthe deep­
this pointin each
of sampling, the

boat was tied to the buoy, obviating the necessity of dropping anchor
each time.
For purposes of sampling, four levels were selected.
(1)

These were;

central epilimnion, taken always at three feet? (2) top of thermo-

cline——this depth varied, but was established at each sampling time
with a vertical series of temperatures; the mean depths for this level
for each of the lakes were, Starvation——7.4 feet, Timijon——9,1 feet,
Juanita — 7.7 feet, Irwin— 11.5 feet, and Grant1s— 16.5 feet; (3) cen­
tral hypolimnion, usually 20 feet; and ^4) bottom waters, one foot
above the mud-water interface.

O *H

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17
Chemical sampling
Water samples were collected from the four levels described above
with a modified Kemmerer water sampler and analyzed immediately for
dissolved oxygen by the rapid Winkler method (Ellis, Westfall, and
Ellis, 1948) with the Alsterberg (sodium azide) modification ("Stan­
dard Methods for the examination of water,

sewage, industrial wastes,"

American Public Health Association, 1955); for free carbon dioxide by
the method given by Ellis, Westfall, and Ellis (1948); for alkalinity
according to "Standard Methods"; for pH with one of the following:
Hellige pH comparator, Coleman Compax pH meter, or Beckman pH meter,
Model N; and for color with a USG-S color kit using standard glass discs
calibrated according to the platinum-cobalt color scale.

Water tempera

tures were taken with a pocket thermometer immediately after the sample
was collected with the sampler; air and surface temperatures were taken
directly.

Light penetration was measured with a Secchi disk.

In addi­

tion to the above determinations made in the field, water samples were
collected from all four levels and bottled for return to the field
laboratory where they were analyzed for total hardness by the EDTA
(versenate) method ("Standard Methods") and for conductivity with a
conductivity bridge manufactured by Industrial Instruments, Inc.

Simi­

lar water samples were also collected (excluding 1953) and stored for
later determination of total phosphorus, when the method of Ellis,
Westfall, and Ellis (1948) was used with a Klett-Summerson photo­
electric colorimeter.
In addition to the above analyses made of the water, mud samples
were collected from the two treated lakes, Starvation and Timijon, to
be analyzed for adsorbed calcium in an attempt to determine if intro­
duced calcium from the applied lime was being lost due to ion-exchange
at the mud-water interface.

The samples were collected always at the

18
sampling station with an Ekman dredge, making sure that the interface
was held within the dredge.

Approximately the surface three centimeters

were scooped out, filtered through filter paper (Whatman No. 1), and
air-dried for storage.

For determination of adsorbed calcium, the

method of Cheng and Bray (1951) was used with modifications.

The sam­

ples were oven-dried at 60°C. and ground to a 200-mesh size.

To one

gram of this material was added 10 ml. of 23% NaNO^ as the extracting
solution; this mixture, after shaking for one minute and standing for
ten minutes, was filtered through filter paper (Whatman No. 1) and the
filtrate was analyzed for total hardness (calcium and magnesium) by the
EDTA (versenate) method (’'Standard Methods").
Because some of the mud samples collected after alkalization constained free carbonates (fall-out from the autumn application), the
method of Hissink (1923) was modified and utilized.

Hissink’s method

consisted of carrying out two leachings upon each sample, assuming that
all the adsorbed bases would be removed by the first leaching and that
equal amounts of bases would be removed through dissolution of the car­
bonates by the extracting solution in the two leachings; the second
leaching was then subtracted from the first in order to obtain the ad­
sorbed bases.

In the samples of the present study, three leachings

were made to test the above assumptions (i.e., if the assumptions
were valid, the results of the second and third leachings should have
been equal) which were found to be invalid.

Therefore, calculations

of adsorbed calcium were made in the following manner:

It was still

assumed tnat all adsorbed calcium was removed in the first leaching,
but it appeared that greater amounts of bases, derived from free car­
bonates, were removed by prior leachings.

The third leaching was sub­

tracted from the second, and the difference was added to the second;

19
this sum was then subtracted from the first leaching, the difference
being an estimate of adsorbed calcium.

Biological sampling
Two samples of net plankton were collected from each of the four
levels or each sampling date with a Juday plankton trap holding 10
liters, with a net made of No. 20 silk bolting cloth.

The contents

were stored in one—ounce bottles in a resulting preservative of about
3 or 4% formalin.

In the laboratory, all samples were brought up to

a uniform volume of 30 ml.

The samples from the two hypolimnial levels

were found to contain negligible amounts of plankton (other than Chaoborus) and were not included in the plankton analysis.

The samples

from the two epilimnial levels were analyzed for volume of net phyto­
plankton and zooplankton.

The method of Dice and Laraas (1936) for

graphically comparing several sets of data was employed for all biolo­
gical data*

The standard error of the estimate was computed for each

pair of duplicate samples; two standard errors were plotted on each
side of the mean as a straight vertical line.

In comparing any two

sampling dates, if the vertical lines did not overlap horizontally,
statistical significance was indicated*
Net phytoplankton
For the determination of volume of net phytoplankton, a SedgewickRafter counting cell was filled with one ml • of the concentrate, and
under the microscope, a total of 10 fields per sample were counted,
using a Whipple ocular micrometer; an average volume per cell was cal­
culated for each species, using measurements made with a calibrated
ocular micrometer and approximating the shape of the cell with some
geometric figure (method modified from Whipple, 1948, p. 119).
the counts and average volumes,

From

standing crops in terms of volume per

20
liter were calculated*

Zooplankton
To determine the volume of zooplankton per sample, a different
method was used on each of the following two groups comprising the
zooplanktons

(1) smaller organisms, composed of rotifiers and nauplii,

and (2) larger organisms, composed of Cladocera and Copepoda.

For the

rotifers, a strip of concentrate in the Sedgewick-Rafter cell one milimeter wide and 50 milimeters long was observed and all rotifers and
nauplii were counted*

For the large crustaceans, the entire area of

the Sedgewick-Rafter cell was observed and all cladocerans and copepods
were counted*

For both groups, an average volume was calculated from

measurements for each species, and from counts and these average volumes,
the standing crops, as volume per liter, were computed*

RESULTS AND DISCUSSION
Preliminary laboratory experiments
Release of phosphorus
To test for the release of phosphorus, hydrated lime was added to
bottles containing water and mud, and after three days, the water was
analyzed for dissolved phosphorus.

Three mud-water combinations from

each of the three colored bog lakes were treated with lime: (1) surface
water with mud,

(2) bottom water with mud, and (3) surface water with­

out mud (control).

Six different rates of lime application were used:

0 (control), 25, 50, 100, 150, and 200 pounds per acre-foot.

In all

cases, the experimental unit consisted of 50 ml* of mud, taken from
the mud-water interface, added at the bottom of one liter of water in
a two-quart jar.

The lime was added by dissolving previously weighed

amounts of lime with water from the experimental bottles and adding

21
the solution to the bottle.

Water samples were extracted for analysis

by pipette three days after the lime addition.

Dissolved phosphorus

was determined by the method given by Ellis, Westfall, and Ellis (1948),
using a Klett-Summerson photoelectric colorimeter.
The results are shown in Figure 6, where it can be seen that
dissolved phosphorus increased in the series which included mud, with
one exception, when the lime application rates increased.

The excep­

tion, Starvation Lake bottom water, which had an original high concen­
tration of dissolved phosphorus,
lime application.

showed no increase of phosphorus with

In all control bottles (no mud) there was no increase

of dissolved phosphorus with lime application, indicating that the re­
leased phosphorus in the other bottles did, in fact, come from the mud.
The release of phosphorus upon alkalinization may be due either
to (1) raising the pH to a more suitable level for decomposition bac­
teria, thus mineralizing phosphorus fixed in an organic form, or to
(2) an anionic exchange between adsorbed phosphate and introduced hy­
droxide.

No data are shown here to indicate that either of the above

phenomena was in operation to the exclusion of the other.
Calcium adsorption
To test for calcium adsorption, experiments were set up with bottles
containing water and mud which were treated with hydrated lime; perio­
dical tests for alkalinity and total hardness were made, and finally
a solution of sodium nitrate was applied and the water subsequently
tested for calcium.
Three mud-water combinations were used from each of the three
colored bog lakes, the amount of water being one liter in all cases
and the amounts of mud being 0 (control), 50, and 100 ml.

Hydrated

lime was applied at the rates of 50 pounds per acre-foot for Juanita

22

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23
and Starvation lakes, and 100 pounds per acre-foot Tor Timijon Lake*
Hydrated lime was added to the bothies

in a manner similar to that

described above for the phosphorus experiments*

Tests for alkalinity

and total hardness were made periodically for about five months, at which
time the sodium nitrate solution was added to all bottles at the rate
of ten times the equivalents of the calcium hydroxide (hydrated lime)
originally added*

Tests for alkalinity and total hardness were again

made 10 days after the addition of sodium nitrate*

Alkalinity was de­

termined by methods given in "Standard Methods” pmd total hardness by
the EDTA (versenate) method ("Standard Methods")*

(The total hardness

test is a determination for both calcium and magnesium, but since the
hydrated lime used contained practically no magnesium, this report is
limited to the discussion of calcium*)
The results show that calcium was lost from the water after lime
application, as smaller concentrations of total hardness were observed
in the bottles containing mud, as shown in Figures 7, 8, and 9*

Fur­

thermore, concentrations of total hardness were smaller in the bottles
containing the greater amount of mud, indicating that more than the
mud-water interface was involved in cation-exchange phenomena.

Barrett

(1952) after conducting similar bottle experiments with mud and water
from another lake, also suggested that more than the interface is active
in calcium adsorption*

Alkalinity was found to be lower in those bot-

les containing mud than in those without mud*
■When the sodium nitrate solution was added, increases of dissolved
calcium were observed in all bottles containing mud, indicating that the
calcium which had been introduced and adsorbed by the mud was released
through cation exchange between calcium and sodium.

Alkalinities did

not increase to high values after the addition of sodium nitrate because
the calcium, after release, was probably in the form of calcium nitrate,

24

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27
which does not contribute bo alkalinity*
It should be pointed out that the total hardness of the water in
the bottles which contained mud increased upon application of sodium
nitrate to values above those observed in the control bottles without
mud; this increase was due to a greater concentration of released cal­
cium becau se the volume of water had been reduced through the time
of the experiment by withdrawing samples for analysis.
fore, do not show exact quantitative relationships.

The data, there­

It was concluded,

however, that adsorption of introduced calcium did occur, and that ad­
sorption was greater in the presence of a greater amount of mud.
Field experiments
The immediate effects of the lime application were to raise the
pH of the lake water, and to increase the alkalinity, total hardness,
and conductivity.

The summer application— while stratification was

present-resulted in chemical effects in the epilimnion only; no ef­
fects were observed below the thermocline *

The autumn application—

when the lakes were nearly homothermous— resulted in almost the entire
application of the lime, in dissolved form, falling to the bottom levels
of the lakes, apparently because of the high density of the lime-water
discharge solution; the dissolved lime remained at the bottom for the
duration of the experiment (through the following winter and summer)
and did not become dispersed throughout the upper levels of water.
Plankton responses were observed in Starvation Lake soon after the lime
treatment, and in both treated lakes the following summer.

Conditions

in the control lake, Juanita, remained essentially constant during the
time of the experiment.
Tables containing complete data on the field experiments are to
be found in the appendix; graphs and abbreviated tables illustrating
the more important effects accompany the following detailed discussions.

28
Starvation Lake
Effects in epilimnion*

The summer application to this lake was

na de as rapidly as possible and brought about large and precipitous
changes in the chemistry of the lake waters; because of the strong
thermal stratification, the effects of the treatment were felt only in
the epilimnion*

The pH rose from approximately 5*5 to almost 11*0 in

the central epilimnion, and then descended more slowly to level off at
about neutrality in about 5 weeks, where it remained for the rest of the
summer (see Figure 10)*

The autumn application had little if any effect

upon the pH in the epilimnion, and through the summer of 1955 the pH
here remained about neutral*

In Figure 10 are also shown the data con­

cerning alkalinity, and it can be seen that these data follow what would
be expected with the corresponding pH values*

Total alkalinity increased

from about 4 ppm to 70 ppm in the central epilimnion; some of the alka­
linity was in the form of hydroxide immediately after lime application,
but this soon changed to normal carbonate and eventually to bicarbonate
as carbon dioxide was produced by the respiratory processes of the lake
and was made available for combination with the hydroxide and carbonate*
Dissolved carbon dioxide was not present, of course, immediately after
the application, because of its rapid utilization in combining with the
introduced hydroxide; free carbon dioxide appeared again when the pH
values descended to about 8*0*

Dissolved oxygen concentrations did not

appear to be affected directly by the lime application*

Figure 11 shows

the changes in total hardness through both years for the central epi­
limnion, and it can be seen here that the total hardness generally fol­
lowed the same trends as did total alkalinity*

Conductivity, though

not in the same units, followed the same general trends as did alkalinity
and total hardness (see Figure 11)#
The effect of alkalinization upon color in the epilimnion was parti-

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31
cularly observed in Starvation Lake in order to test the postulate
which Hasler et a l ♦ (1951) supported experimentally in Wisconsin, namely,
that the color would be precipitated*

In Starvation Lake, no decrease

in color was observed following the lime application; rather, an increase
was observed shortly after the application, and later, the high values
fell off to approximately the same as before lime application (see Fig­
ure 12)•

As would be expected, Secchi disk readings were reduced at

this time of apparent higher color concentration (see Figure 12)*

Dur­

ing this time, it was observed in water samples taken in conjunction
with chemical analyses made in the field, that strands of brown, mucilaginous-appearing material were present in the water, particularly
at the top of thermocline level, where the increase in observed color
was even greater than in the central epilimnion.

Following the theory

of color flocculation and precipitation by calcium, it is suggested
that a partial flocculation did take place, producing the brown strands
in the water; some precipitation took place also, and a concentration
of the material accumulated on top of the thermocline, probably because
of the increased water density at that level.

Apparently this partial

flocculation, with the resulting visible brown strands, effected an
apparent increase in color when measured with the USGS color kit.

The

color at all levels returned to normal values after about four weeks.
Chemical conditions at the top of thermocline, generally followed
trends similar to those in thecentral epilimnion, except that changes
were not as great, and concentrations of dissolved salts did not increase
to the high values observed in the central epilimnion.
Effects in hypolimnion.

Conditions in the central hypolimnion,

and in the bottom waters, did not change until after the autumn appli­
cation.

At the time of the autumn application in October, water tempera-

32

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33
tures varied from 49°F. at the surface to 41°F* at the bottom, with
no thermocline present.
upon liming.

Conditions in the epilimnion did not change

Conditions in the central hypolimnion changed slightly but

significantly, while changes in the bottom waters were extremely pre­
cipitous, being the result of almost the entire lime application fall­
ing to the bottom in solution.

pH values rose from about 6.0 to 11.0,

total hardness from 11 to 118 ppm., total alkalinity from 17 to 128 ppm.,
and conductivity from 19 to 147 micromhos.
The high density of the solution at the. bottom following this
autumn application apparently mode possible a temperature inversion
and the formation of a monimolimnion.

Temperatures at the bottom

were increased slightly (even above temperatures found in the next
higher level) from 41°F. before alkalization to 44°F. after alkalization; dissolved oxygen also increased at this bottom level from 0.0
to 2.0 ppm. after liming.

These changes were brought about by the use

of surface water— which was warmer and contained dissolved oxygen—
for mixing with the lime in the mixing tank on the raft.

The lime-

water suspension, containing oxygen and at a higher temperature, was
more dense than the coolest water in the lake at the bottom; therefore,
when it was discharged into the surface of the lake it immediately fell
with little mixing to the bottom levels.
Effects upon phosphorus concentration.

As mentioned previously

in connection with the laboratory experiments with mud and water in
bottles, the phosphorus concentration of the bottom waters in Starva­
tion Lake was high, even before any lime application, and no apparent
changes were brought about upon treatment in the bottle.

The same re­

sults were observed in the lake after the autumn lime application was
made.

The mean total phosphorus concentration in the bottom waters

before the autumn application (1954 only) was 214 ppb., while the mean

34
after the autumn application (1955) was 244 ppb#; a t-test showed a
t-value of 1,88 with 16 degrees of freedom, the t-value not being
significanti
The total phosphorus concentration in the central epilimnion appar­
ently increased immediately after the summer application, as shown in
Figure 13#

(These data, however, are not conclusive; other apparent

increases in total phosphorus were observed at different times and levels,
and in other lakes, suggesting variation due to causes other than the
lime treatment*)

The source of released phosphorus was probably-in

those shallower areas of the lake where epilimnial waters, containing
low concentrations of phosphorus,

came into contact with bottom muds;

these conditions are essentially those produced in the bottle experiments
with "mud and surface water", where the release of phosphorus was ob­
served#
It might be suggested that any observed increase in phosphorus
in the epilimnion was due to phosphorus actually present in the liming
material used#

However, this was not the case, as an analysis of the

hydrated lime showed only a negligible amount of phosphorus present#
A lime solution containing 75 ppm. of alkalinity was found to contain
only about one ppb# phosphorus (well within the error of the analytical
method), while the increase in phosphorus observed in the lake (for an
increase of about 75 ppm# alkalinity) was about 40 to 50 ppb#

It was

concluded, therefore, that there were no increases in phosphorus due
to introduction with the lime.
Calcium adsorption by m u d *

The summer application of lime had

no effect upon the adsorbed calcium in the pulpy peat; this vo uld be
expected, considering that the summer application did not affect chemi­
cal conditions in the bottom waters#

The autumn application, however,

35

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36
resulted in alkalinization of the bottom waters only, and the mud
samples taken after the autumn lime application showed an increase in
adsorbed calcium.

As per cent calcium on an oven-dried basis, adsorbed

calcium was 0*26% before the autumn application (1953 and 1954) and
0*38% after the autumn application (1955)*

A t-test showed a t-value

of 7.74 with 36 degrees of freedom, the t—value being highly signi­
ficant.
The modification of the scheme of Hissink* s (1923) which was used
for determining adsorbed calcium in the presence of free calcium car­
bonates precludes the determination of absolute values of either the
percent figures quoted or the degree of increase.

In the Starvation

Lake mud samples, free carbonates could not be seen (in Timijon Lake,
as will be pointed out later, carbonate material could easily be seen
in the samples), and that serious interference from free carbonates
was encountered is doubtful.

The data can probably be accepted to the

degree that a significant increase of adsorbed calcium occurred, but
exact quantitative values are not implied.
Biological effects.

In the analysis of plankton samples, phyto-

plankton and zooplankton were separated on the presumption that the two
groups belonged to different trophic levels.

The reason for using dif­

ferent analytical methods for determining the total volume per sample
was that in the latter group, zooplankton, the organisms were large but
relatively scarce, while in the former group, phytoplankton, the oppo­
site situation prevailed, and a common analytical method would result
in extreme variation.

Hven among the zooplankton, two methods of analy­

sis were necessary because of the presence of rotifers, which were small
but more plentiful in relation to the larger copepods and cladocerans.
The different analytical methods were employed in order to reduce the

37
variation which would result from using, for example, a method designed
to sample small, abundant organisms, such as phytoplankton, upon larger,
less abundant organisms,

such as zooplankton.

Even with this system, a major difficulty presented itself in the
presence of a phytoplankter, Peridinium liinbatum, which was very large
in comparison with other phytoplankters, and which caused an extreme
variation in the analytical results*

For this reason, data concerning

Peridinium limbatum were removed from the volume computations in con­
structing the graphs of standing crop of net phytoplankton for all lakes*
Data on Peridinium limbatum are included, however, in the tabulated re­
sults given in the appendix.
The standing crops of net phytoplankton for the central epilimnion
are given in Figure 14*

It will be noted that net phytoplankton abundance

was extremely low during that part of 1953 sampled, and for all of 1954*
A decrease in phytoplankton was observed immediately after application,
apparently being the result of inhibitory effects of the lime suffered
by the organisms; Whipple (1948, p. 405) discusses the use of hydrated
lime as an algicide and says that lethal effects are probably due to the
loss of carbon dioxide available for photosynthesis or to changes in
hydrogen ion concentration.

During 1955, 'however, large, statistically

significant increases in standing crop were observed*

The principal

organism making up this "bloom" was Dinobryon sertularia.
During 1954, shortly after the summer lime application, a bloom of
Microcystis aeruginosa occurred in the epilimnion; this bloom is not shown
among the plankton data because the cells were of nannoplankton size and
were not collected in the plankton samples, being too small to be re­
tained by the plankton net (No. 20 silk bolting cloth).

The bloom was,

however, obvious to the eye; it gave a definite greenish color to the
water, and collected in windrows of "scum" along the lee shores of the

38

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39
lake at the edge of the bog mat.

The date when it was most strikingly

observed was August 2, 1954, and chemical data for this same date re­
flected the effects of the bloom.

Dissolved oxygen increased markedly

from the previous sampling date, from 2,8 to 10,5 ppm,; a rise in pH
was observed with a corresponding shift in the form of alkalinity from
the bicarbonate form to the normal carbonate form, results of carbon
dioxide extraction from the bicarbonate (see Figure 10); and the Secchi
disk reading was markedly decreased (see Figure 12),

The bloom occurred

while the phosphorus content of the epilimnion was high (after the re­
lease of phosphorus discussed previously) which decreased after the
commencement of the bloom (see Figure 13),
Immediately upon lime application the standing crop of zooplankton
was markedly reduced, also apparently suffering inhibitory effects from
the lime treatment.

During the latter part of 1954, however, the zoo­

plankton appeared to have recovered and an increase was observed, when
the standing crop reached levels much higher than observed previously.
This may have been due, in part, to the available food supply produced
by the bloom of Microcystis aeruginosa.

During 1955 the standing crop

of zooplankton appears to have been high, remaining above pre-alkalization levels through the 1955 sampling season, although statistical signi­
ficance is not shown for all sampling dates in 1955,

Standing crops of

zooplankton are shown in Figure 15,
A complete list of the species of the plankton found in Starva­
tion Lake is given in Table 1.
Timijon Lake
Effects in epilimnion.

The summer lime application in Timijon Lake

was made over most of the summer of 1954 and at such a rate as to keep
the pH of the epilimnial waters approximately at neutrality; this was

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Table 1.

List of plankton organisms, Starvation Lake*

Phytoplankton
Chlorophyta
Pietyosphaerium pulchellum Wood
Dimor phococcus lunatus A* Braun
Pandorina sp^
Desmidiaceae
Desmidium sp*
Staura strum sp*
Xanthidium sp*
Chrysophyta
Dinobryon sertularia Ehrenberg
Bacillariophyceae (diatoms)
Asterionella sp *
Cyanophyta
Miorocystis aeruginosa Kuetz*
Euglenophyta
Euglena sp*
Pyrrhophyta
Glenodinium sp•
Peridinium limbatum (Stokes) Lemmermaim

Zooplankton
Rotifera
Anuraea cochlearis Gosse
var. maoracantha
Notholca longispina Kellicott
Polyarthra platyptera Ehrenberg
("others uni dent if i o d )
Cladocera
Bosmina sp*
Daphnia sp*
Holopedilum ^ibberum Zaddach
Copepoda
Cyclops sp.
Dia-ptomus sp •

42
done in an attempt to avoid "the killing of plank "ton which had been
observed in Starvation Lake*

As a result, the pH values never reached

such high values in Timijon Lake as in Starvation, nor did alkalinity
reach such high concentrations as observed immediately after alkalization in Starvation, although alkalinity values were about the same in
the two lakes at the end of the summer season.
At the first small lime application in Timijon, the pH rose from
5*3 to 6*7 in the central epilimnion; subsequent small applications were
made as the pH fell below neutrality due to combination with carbon diox­
ide, each small application usually resulting in a rise of pH to between
7 and 8,

At the close of the summer sampling period, the pH in the cen­

tral epilimnion was 8*5, and 7*4 in October immediately before the autumn
lime application was made*

The autumn lime applic ation had only a slight

effect on the pH in the central epilimnion, but did have a great effect
at the top of thermocline, as well as in both hyplimnial levels*

pH

values during 1955 remained at about neutrality for the duration of the
summer*

pH values are shown in Figure 16; for sampling dates upon which

lime applications also were made, the data given are those obtained
before the applications, rather than after*
Changes in alkalinity are also shown in Figure 16*

During the time

of lime application in the summer, the total alkalinity increased by jumps
as each small application was made, and in October, before the autumn
application, total alkalinity was 23 ppm*

The autumn application in­

creased the alkalinity slightly in the central epilimnion, but by a
greater amount at the top of thermocline.

No hydroxide alkalinity was

ever observed in the epilimnial levels, although normal carbonate was
found, usually immediately after a lime application.
hardness and conductivity,
those of total alkalinity*

Changes in total

shown in Figure 17, followed, in general,

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45
Figure 18 shows "the changes in color in both epilimnial levels.
The color observed in the central epilimnion did not increase abruptly
during or after the summer lime application, as it did in Starvation
Lake, nor did it increase after the autumn application.

At the top of

thermocline, however, where a greater increase in alkalinity and pH
values were observed immediately after the autumn lime application,
color increased abruptly and significantly, much as it did in Starvation
Lake; Secchi disk readings were reduced from 9 feet to 6 feet, the low­
est ever observed in Timijon Lake.

This apparent increase in color and

also that observed in Starvation Lake appear to be the result of a large,
sudden change in alkalinity with accompanying high pH values, rather
than of total change of alkalinity.
Color appeared to have undergone a decrease in both epilimnial
levels during 1955.

This was probably due to the small amount of precipi­

tation received during the summer of 1955, which reduced the run-off
water carrying allochthonous material responsible for color into the
lake.
Effects in hypolimnion.

The summer application of lime had no

effects upon either of the two hypolimnial levels.

However, at the

autumn application in October, 1954, when the lake was nearly homothermous (temperatures ranged from 50°F. at the surface to 42°F. at the bot­
tom), the same event occurred as in Starvation Lake; that is, the dis­
solved lime fell to the bottom, in solution, increasing the alkalinity,
pH, total hardness and conductivity at both hypolimnial levels.

(As

reported above, some effects were also felt in the epilimnion.)

pH

■values rose in the bottom waters from about 6 to 11, total alkalinity
increased from 13 to 175 ppm. (including a large proportion of hydroxide
alkalinity), total hardness from 11 to 154 ppm, and conductivity from

46
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47
16 "to 286 micromhos.

As in Starvation Lake, a temperature inversion

appeared after lime application; "temperatures in the bottom waters
rose from 42°F. to 46°F,, and dissolved oxygen increased from 0 to
6 ppm*

Effects upon phosphorus concentration.

No changes in total phos­

phorus concentration were observed in the epilimnion during or following
the summer lime application.

In the bottom xvaters, however, total phos­

phorus increased after the autumn lime application from a mean during
1954 of 23 ppb, to a mean during 1955 of 156 ppb.; a t-test showed a
t-value of 10*99 with 16 degrees of freedom, the t-value being highly
significant.

This result was essentially the same as obtained in the

laboratory bottle experiments with "mud and bottom water," where the
release of phosphorus was observed.
It might be argued that this large increase observed in the bottom
waters might not have been due to a release of phosphorus caused by
liming.

It is still in doubt whether the meroinictic conditions which

appeared to be so effective in the bottom waters over the winter of
1954-1955 (the high concentration of alkalinity in the bottom waters
was not dispersed) were natural conditions or whether the meromixis was
induced by the introduction of the layer of dense (alkaline) solution.
If the latter case were true, the factors which generally are respon­
sible for a high concentration of phosphorus at the bottom of meromictic lakes might also have been effective in producing the high concen­
trations observed in this case.
However,

certain evidence points to the conclusion that Timijon Lake

was meromictic under natural conditions.

No analyses of water from the

bottom waters ever showed a trace of dissolved oxygen.

In the central

hypolimnion, which was usually 20 feet above the bottom, oxygen determina­
tions showed only small concentrations, if any, and then only in the early

48
spring*

Furthermore, when chemical analyses were made in October, 1954,

just previous to the autumn lime application, thermal stratification
had been destroyed, but dissolved oxygen was absent at both hypolimnial
levels*
Finally, the results of the bottle experiments which showed re­
leases of phosphorus from the mud lend supporting evidence to the postu­
late that liming would release phosphorus in the lake*
Calcium adsorption by m u d *

The summer lime application had no

effects upon the adsorbed calcium in the pulpy peat at the mud—water
interface*

However, adsorbed calcium, as percent calcium on an oven-

dried soil basis, increased from 0*24% before the autumn application
(1953 and 1954) to 0*34^ after liming (1955)*

A t-test showed a t-value

of 4*81, degrees of freedom being 36, the t-value being highly signifi­
cant*
Here again, as explained also in the case of Starvation Lake, the
assumptions of H i s s i n g s method were not met in the analytical technique*
Free carbonates were definitely present and plainly visible to the eye,
and probably interfered considerably with the accuracy of results*

It

is probably safe to conclude that adsorbed calcium did increase, but
such a conclusion must be made primarily upon the basis of there being
present greater concentrations of calcium available for cation exchange
and upon the indirect evidence shown in the results of the bottle experi­
ments which showed definite exchanges of calcium.
Biological effects.

In Timijon Lake the same difficulty was en­

countered with the phytoplankter, peridinium limbatum, as in Starvation
Lake*

For the reasons expressed previously in connection with Starva­

tion Lake, this organism was excluded from the data in constructing the
graphs but is included among the tabulated data in the appendix*

49
Figure 19 shows the changes in standing crop of net phytoplankton
in the central epilimnion.

It can be seen that net phytoplankton was

scarce for all sampling dates during 1953 and 1954, but that increases
occurred during 1955, although these increases do not approach the very
high values found in Starvo.tion Lake.

The principal organism making

up the large standing crop observed in 1955 was Dinobryon sertularia,
which was the same in the case of Starvation Lake.
Figure 20 shows the changes in standing crop of zooplankton in the
central epilimnion.

Among the samples of this group, more variation

was observed than in net phytoplankton, but in general, the volume of
zooplankton was higher in 1955 than during 1953 and 1954.

As was the

case with Starvation Lake, the zooplankton in Timijon appeared to suf­
fer inhibitory effects upon the first lime application during the summer
of 1954; a sharp decrease was noted in zooplankton immediately after the
first application, and no recovery was observed during the latter part
of the 1954 summer (which was observed in Starvation Lake), the probable
reason being that, in the case of Timijon Lake, lime applications were
continued through the 1954 summer which probably had a continuing in­
hibitory effect.
All species of phytoplankton and zooplankton are listed in Table 2.
Juanita Lake
Chemical conditions.

Generally, the observation made on this lake

were constant over the period of the experiment.

Table 3 shows mean

values for each of the four levels sampled for color, pH, bicarbonate
alkalinity, total hardness, conductivity, and total phosphorus, and also
for light penetration (Secchi disk reading), averaged over the entire
time of sampling (August, 1953 through 1955); the detailed data are pre­
sented in tabular form in the appendix.
As this lake was the control for the lime application experiments

Indicates sampling date, volume less than O.Ol mm3

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52
Table 2#

List of plankton organisms, Timijon Lake*

Phytoplankton
Chlorophyta
Dictyosphaerium pulchellum Wood.
Pandorina sp •
Desmidiaceae
Staurastrum sp#
Chrysophyta
Dinobryon sertularia Ehrenberg
Synura uvella Shrenberg
Bacillariophyoeae (diatoms)
Asterionella sp#
Tabellaria fenestrata (Lyngb#) Kuetz#
Eugleno phyta
Euglena sp•
Pyrrhophyta
Glenodinium sp#
Peridinium limbatum (Stokes) Lemmermann

Zooplankton
Rotifera
^nuraea aculeata Gosse
Anuraea cochlearis Gosse
v a r • macracantha
Notholca longispina Kellioott
Polvarthra platyptera Ehrenberg
Cladocera
Bosmina sp»
Da-phnia sp.
Holopedilum gjbberum Zaddach
Copepoda
Cyclops sp•

53

Table 3. Mean values of color, pH, alkalinity, total hardness,
conductivity, total phosohorus, and light penetration,
Juanita Lake*

Level

Central
Epilimnion

Color

90

120

5.0

4.9

4.8

5.1

Alkalinity
(bicarbonate)
ppm# CaCO-j

4

4

3

6

Total
Hardness
ppm. CaCO^

5

5

5

5

7

7

8

8

10

14

16

72

pH

Top of
Thermocline

Central
Hypolimnion

Bottom
Waters

115

150

Conductivity
mho X 10“6

Total
Phosphorus
ppb. p

Light
penetration

(Secchi disk reading, feet)

7

54
carried out on Starvation and Timijon lakes, it was sampled on a schedule
similar "to "those lor "the "treated lakes, and some points should be pointed
out as being pertinent.

Dissolved oxygen levels remained usually between

4 and 6 ppm., except lor some samples in the spring, when cooler tempera­
tures prevailed and oxygen concentrations were somewhat higher; this
was probably due to a greater solubility at these cooler temperatures,
greater absorption ol oxygen from the air because of greater wind-induced
turbulence, and to the lact that decomposition processes, more active
at warmer temperatures, had not yet consumed dissolved oxygen to a
large degree.

The amount ol color in the water appeared in a strati­

fied manner; that is, less color was present at the shallower levels.
A slight increase in pH was found in the bottom waters, while alkalinity
also was at a slightly greater concentration at this level.

Usually,

oxygen was absent from both hypolimnial levels, but occasionally occurred
in the central hypolimnion in the early spring in low concentrations,
indicating that partial spring circulation had been effective in bringing
dissolved oxygen to at least below the level of the thermocline.

Dissolved

carbon dioxide was always present at all levels, reaching higher concen­
trations at the deeper levels (it should be pointed out that carbon diox­
ide determinations cannot be accepted as completely accurate in waters
which contain high amounts of colloidal humic materials, which act as
humic acids and which may be titrated along with carbonic acid in the
carbon dioxide test).

Neither hydroxide nor normal carbonate alkalinity

was ever observed.
Some differences in total phosphorus concentration were observed
in the bottom waters, as was the case in Timijon Lake, but the statisti­
cal significance of the difference between the values observed in 1954
and those observed in 1955 is doubtful.

Mean total phosphorus in the

bottom waters for 1954 was 64 ppb., and for 1955 at the same level was

80 ppb.

A t-test showed a t-value of 2.60 for 15 degrees of freedom,

the t-value being significant at the b% level only.
Chemical conditions in the two treated lakes were generally the
same as those observed in Juanita Lake except for changes brought
about by lime application, and it can be concluded that lime applica­
tion was responsible for chemical conditions that deviated from those
observed in Juanita Lake*

Biological conditions.

Figure 21 shows the standing crop of net

phytoplankton in the central epilimnion for the entire sampling period.
It will be seen that, in general, the standing crop was low and, during
1953 and 1954, was approximately the same as in Timijon and Starvation
lakes; during 1955, the standing crop in Juanita Lake was definitely
less than in the two treated lakes.

It appears that there may have been

some increase in net phytoplankton during 1955, but most sampling dates
do not show statistically significant differences, except at the close
of the summer sampling period.

In Juanita Lake, the organisms which

accounted for "the bloom in the two treated lakes, namely, Microcystis
aeruginosa and Dinobryon sertularia, did not appear at all.

The data

which were used for constructing the graphs, as in the cases of Timijon
and Starvation lakes, do not include Peridinium limbatum, but data on
peridinium. limbatum are included in the tables in the appendix.
If the data concerning Peridinium limbatum were included in the
construction of the graphs in Figure 21, an increase would be apparent
at the close of the 1955 sampling season, particularly on the last
sampling date.

No explanation of this increase is offered here, but

the significance of such an apparent increase is doubtful in the light
of the variation caused by this organism*
Figure 22 shows the changes in standing crop of zooplankton in
the central epilimnion, and it can be seen that some apparent increases

56

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58
occurred during 1955*

Not all "the differences, however, were statist!—

cally significant when compared "to earlier data, and were not of as
grea"t a magnitude as the changes which occurred in the two treated lakes•
Table 4 lists by species all plankton organisms found,
Irwin Lake
Samples were obtained from Irwin Lake and Grant’s Lake, both of which
were clear—water bog lakes, for the purpose of making comparisons with
the two treated colored lakes, particularly in the event that the lime
treatment cleared the color from the treated lakes.

The fact that the

colored lakes were not cleared upon alkalization precludes a comparison
on this basis*

However, data on the two clear lakes are included here,

and comparisons are made concerning the differences in chemical and bio­
logical conditions*
Chemical conditions *

Chemical analyses showed that bicarbonate

alkalinity, total hardness, conductivity, and pH were approximately
the same in Irwin Lake as in any of the colored lakes before lime treat­
ment*

Light penetration (Secchi disk reading) was deeper and color con­

centration less than in the colored lakes*

Dissolved oxygen relation­

ships were entirely different in Irwin Lake, in that oxygen was ordin­
arily found in depths well below the thermocline, even in the bottom
waters during times of thermal stratification, and also in that concen­
trations of oxygen in the epilimnion were generally higher than in the
colored lakes*

Carbon dioxide concentrations were found to be less at

the deeper levels than in the colored lakes, but this may be due, in
part, to the absence of humic acids and, in part, to a greater amount
of photosynthesis being carried out at these depths.

The greater pho­

tosynthesis, made possible by the penetration of light througn the clearer
water to below the thermocline, also accounts for the presence of dis-

59
Table 4*

List of plankton organisms, Juanita Lake

phyto p 1ankton
Chlorophyta
Dictyosphaerium pulchellum Wood
Desmidi&ceae
Staurastrum sp*
Xanbhidiuiii sp•
Chrysophyta
Dinobryon bavaricum Inhof
Mallomonas sp•
Bacillariophyceae (diatoms)
Asterionella sp*
Tabellaria fenestrata (Lyngb•) Kuetz.
Euglenophyta
Euglena sp•
Pyrrhophyta
Glenodinium sp•
peridinium limbaturn (Stokes) Lemmermann

Zooplankton
Rotifera
“Anuraea coohlearis Gosse
var. maoracantha
Rotholca longispina Kellicott
Polyarthra platyptera Ehrenberg
(others unidentified)
Cladocera
Bosmina sp*
Holopedilmn gibberum Zaddach
Copepoda
Cyclops sp*
Diaptomus sp •

60

solved oxygen below "the "thermocline.

Total phosphorus concentrations

were less than in Juanita Lake, at all four levels.
Only one significant change with time was observed, namely, that
of color, as shown in Figure 23.

Color concentration in the central

epilimnion in August, 1953, was about 4 0 j during 1955, color was reduced
to about 15 or ZOi a similar reduction took place at the top of thermocline.

This change was probably due, as in the case of Timijon Lake,

to less rainfall during 1955 and a reduction of run-off water carrying
colloidal material responsible for the color.
Table 5 shows the mean values of pH, bicarbonate alkalinity, total
hardness, conductivity, and total phosphorus for each of the four levels,
and also Secchi disk readings, for the entire sampling period.

All data

are presented in detail in tabular form in the appendix.
Biological conditions.

Figure 24 shows the standing crop of net

phytoplankton in the central epilimnion over the entire sampling period,
where it can be seen that some increases apparently occurred during 1955.
No changes in the chemistry of lake waters occurred concurrently or pre­
viously to indicate the cause of the increase, although the decrease
of color ^discussed above) which permitted greater light penetration
might have had some causal effect.

A desmid, Staurastrum, accounted

for the higher standing crop because of its large volume, although pre­
sent in small numbers.
Figure 25 shows the standing crop of zooplankton for the central
epilimnion.

No significant changes were observed, but it can be seen

that the standing crop of zooplankton was generally higher than that
found in the colored lakes.
Table 6 lists all species of phytoplankton and zooplankton found
in Irwin Lake.

Color

61

62

Table 5* Mean values of pH, alkalinity, total hardness,
conductivity, total phosphorus, and light penetration,
Irwin Lake*

Central
Epilimnion

Level

Top of
Thermocline

Central
Hypolimnion

Bottom
"Waters

5.2

5.0

5.0

5.3

Alkalinity
(Bicarbonate)
ppm. C&CO 3

3

3

3

4

Total
Hardness
ppm. CaGOg

4

4

4

5

8

7

8

8

5

7

12

49

pH

Conductivity
mho X 10 “ 6

Total
phosphorus
ppb# P

Light
Penetration

.

.

(secchi disk reading, feet)

13.5

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Table 6 .

List of plankton organisms, Irwin Lake,

Phytoplankton
Chlorophyta
Dictyosphaerium pulchellum Wood
Tetraspora lacustris Leiumermann
Desmidiaceae
Staur a st rum sp •
Chrysophyta
Dinobrvon djverii:ens Imhof
UroKlenonsis amerieana (Calkins) Leramerinann
Bacillariophyceae (diatoms)
Asterionella sp,
Tabellaria fenestrata (Lyngb,) Kuetz,
Eu g 1enophyta
sp*
Pyrrhophyta
Glenodinium sp,
Peridinium limbatum (Stokes) Leramermann

Zooplankton
Rotifera
Anuraea coohlearis Gosse
var • inacracantha
Polvarthra platvptera Ehrenberg
(others unidentified)
Cladocera
Bosmina sp,
Dauhnia sp,
fjolopedilum gibberum Zaddach
Copepoda
Cvclops sp,
Diaptomus sp ,

66
Grant1s Lake
Chemical conditions*

Chemical conditions similar to those found

in Irwin Lake were observed in Grant’s Lake.

The pH, alkalinity, total

hardness and conductivity were almost identical in the two clear-water
lakes*

Because Grant’s Lake did not generally become stratified, dis­

solved oxygen and carbon dioxide concentrations usually showed little
differences with depth.

Only three levels were sampled in Grant’s Lake,

because even when stratification was present it was at such a depth
that the level of bottom waters was immediately below the thermocline.
Color concentrations were very low, being even less than in Irwin Lake,
and light penetration was deeper*

Ho significant changes with time

were observed.
Table 7 shows the mean values of color, pH, bicarbonate alkalinity,
total nardness, conductivity, and total phosphorus for each level, and
also of light penetration (Secchi disk readings).

The detailed data

are shown in tabular form in the appendix.
Biological conditions*

Figure 26 shows the standing crop of net

phytoplankton for the central epilimnion.

There were no significant

changes, but in general, the standing crop was similar to that observed
in the colored lakes.
Figure 27 shows the standing crop of zooplankton for the central
epilimnion, where it appears that the zooplankton was more abundant
than in the colored lakes.

At times the standing crop of zooplankton

was much greater at the top of thermocline, indicating a photophobic
reaction of the zooplankton, which was probably maximal in Grant's Lake
because of the extreme clarity of the water.
Table 8 lists all species of phytoplankton and zooplankton found
in Grant's Lake.

67
Table 7• Mean values of color, pH, alkalinity, total hardness,
conductivity, total phosphorus, and light penetration.
Grant1s Lake•

Level

Central
Epilimnion

Color

15

15

25

4*9

5.0

4.8

Alkalinity
(Bicarbonate)
ppm, CaC03

2

3

3

Total
Hardness
ppm# CaC03

4

4

4

7

7

7

3

5

24

PH

Conductivity

Top of
Thermocline

Bottom
Waters

mho X 10"6

Total
phosphorus
ppb. P

Ligftx;
penetration

(Secchi disk reading, feet)

17

68

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70
Table 8.

List of plankton organisms, Grant*s L&ke.

Phytoplankton
Chlorophyta
Desmidiaceae
Staurastrum sp•
Chrysophyta
Dinobrvon sertularia Ehrenberg
Bacillariophyceae (diatoms)
Asterionella sp.
Euglenophyta
Eug;lena sp.

Zooplankton
Rotifera
Anuraea cochlearis Gosse
var, teota
Anuraea cochlearis Gosse
var. macraoantha
Polvarthra platyptera Ehrenberg
(others unidentified)
Cladocera
Bosmina sp.
Daphnia sp.
Holopedilum gibberum Zaddach
Copepoda
Cyclops sp.
Piaptonus sp.

71
GENERAL DISCUSSIONS AND CONCLUSIONS
Effepts of alkalinity upon phytoplankton production
In considering the question of whether a causal relationship
exists between alkalinity and productivity, several concepts must be
kept clearly in mind*
First, if a causal relationship does exist, it may be an indirect
one*

For example, the availability of phosphorus may be greater, caus­

ing higher productivity, under conditions of greater alkalinity; it is
conceivable that increased phosphorus availability might also be pro­
duced under conditions other than those associated with higher alkalinity,
in which case a causal relationship between alkalinity and productivity
could not be inferred*
Secondly, the observed correlation between alkalinity and produc­
tivity appears to be reversed when a certain high level of alkalinity
is reached, resulting in the "alkalitrophy" of lakes; this upper limit
is possibly as low as 120 ppm«, according to Barrett (1953)*

In Con­

necticut, Deevey (1940) reported less production (on the basis of summer
chlorophyll content) in hard-water lakes than in medium-hard-water lakes*
From the standpoint of biological productivity, then, alkalinity can be
too high as well as too low*

In alkalitrophic lakes, very little or­

ganic matter is present in the sediments, and phosphorus, rather than
being present in an adsorbed form (available to plants through ionexchange reactions), is fixed permanently in inorganic precipitates,
probably tricalcium phosphate and associated compounds which are rela­
tively insoluble.

According to Barrett, the fate of phosphorus added

in fertilizers to alkalitrophic lakes is also in inorganic precipitates.
Finally, the reason for increased productivity in more alkaline
waters may be, as suggested by Neess (1948) and Hasler et a l . (1951)
a greater concentration of carbon dioxide available for photosynthesis

72
in "the bicarbonate form#

In this sense, differentiation should be

clearly made between what is being utilized by producing plants and
what is in short supply in the lake; the former is carbon dioxide,
while the latter is calcium*

When lime application is made for the

purpose of increasing the available supply of carbon dioxide, it is
the calcium which is introduced, and not carbon dioxide itself.

The

calcium than acts as a "chemical carrier" (Coker, 1954, p. 45), holding
the carbon dioxide, which has been produced by the respiratory activities
of the lake, in a large "reservoir" of calcium bicarbonate, and the
carbon dioxide then remains available for photosynthesis.

Thus, alka-

lization differs from fertilization, where the material introduced is
utilized directly as a nutrient by plants; in alkalization, only the
"carrier" is introduced.
Free carbon dioxide is not found in high concentrations in surface
waters because it is rapidly lost to the atmosphere upon wind-induced
agitation,

-^uch of the carbon dioxide produced by the respiratory

activities of the lake is lost in this way.

"Water, therefore, is

automatically prodigal of one of its most important biological assets,
and the losses of free carbon dioxide by this means are often great"
(Welch, 1935, p. 95).

One purpose of lime application is to provide

a means of preventing this loss, retaining the carbon dioxide in a
form available for photosynthesis*
To understand the relationships involved in the limitation of
production by carbon dioxide in, for example, a soft—water bog lake,
it would be helpful to know, in addition to the concentrations of free
carbon dioxide and bicarbonates, the rate of carbon dioxide production
by the lake and also the rate of its utilization in photosynthesis
under conditions of observed plankton response, i. e*, a planlrton bloom*
The events which occurred in Starvation Lake in 1954 soon after

73
the summer alkalization afforded the necessary data "to make "the above
calculations and permitted a test "to he made of* "whe"ther "the increased
alkalinity was responsible for increased biological production*
In Figure 10, the graph for bicarbonate alkalinity shows a linear
increase between the dates of July 15, 1954, and July 25, 1954, in the
central epilimnion, covering a total of five sampling dates*

This in­

crease in bicarbonate alkalinity is due to the reaction of calcium car­
bonate (resulting from alkalization) and carbon dioxide (produced by
the lake’s respiratory activities)*

The slope, then, represents the rate

of increase of bicarbonate, and, proportionally, the rate of carbon diox­
ide production.

This rate, calculated by means of the ratios of molecu­

lar weights of the compounds involved, was 1*94 ppm* carbon dioxide per
day.
On August 2, 1954, a phtoplanlcton bloom, caused by a blue-green
alga. Microcystis aeruginosa, was observed with corresponding changes
in water chemistry— higher pH, increased oxygen, decreased bicarbonate,
and less light penetration*

It can therefore be inferred that for the

eight-day period from July 25 to August 2, or for an unknown shorter
period, phytoplankton inoreased rapidly and utilized carbon dioxide
at a rate more rapid than its production*

The utilization of carbon

dioxide could then be calculated*
Table 9 shows the “balance sheet” of carbon dioxide for the eightday period*

Section A is the carbon dioxide available over the eight-

day period, being equal to the sum of that present in the form of bicarbonates and free carbon dioxide at the beginning of the period plus
the carbon dioxide production for the eight days, or a total of 34*9
ppm* carbon dioxide*

The carbon dioxide utilized over the eight-day

period by the phytoplankton,

section B, is equal to that available

(section A) minus that remaining at the end of the period in the form

74

Table 9. Carbon dioxide utilized by phytoplankton after alkalization
compared to carbon dioxide available without alkalization, Starva­
tion Lake (as ppm. COg)*

A«

COg available over 8-day period • • • • • • • • • • • 3 4 «9
Present at beginning of period.
Free C O g ..........................
Half-bound COg (bicarbonate)* • • * 1 9 * 4
Production in 8-day period • • • • • • 15*5

0*0

34.9
B.

COg utilized over 8-day period

......... • • • • • •

22*6

Available over period (section A)* • ■ 34*9
Less COg present at end of period,
Free COg* • • • • • • • • • • • • •
0.0
HaIf-bound COg (bicarbonate)
• • • 12*3

22.6
C»

COg available without alkalization,

.......... . • 20*3

Present at beginning of period,
Free COg* • • • • • • • • • • • • •
3*0
Half-bound COg (bicarbonate). . • • 1,8
Production in 8-day period • . • • • • 15.5
20.3
D«

Portion of COg utilized that was result of lime • • •
COg utilized (section ! ) ) • • • • • • •
Less COo available without
alkalization (section C ) . * * * « *

22*6
20*3
2.3

2.3

75
bicarbonates and free carbon dioxide, or 22*6 ppm. C02 *

In section C,

the carbon dioxide available under conditions of no lime application is
calculated to be 20*3 ppm* CQ^, using the production rate computed above,
and free carbon dioxide and bicarbonate concentrations found before alka­
lization*

In section D is calculated the portion of that carbon dioxide

utilized by the plankton which was the result of alkalization, which is
2*3 ppm* COgj it is the difference between the carbon dioxide utilized
and that which would have been available if no lime application had been
made, for the eight-day period, and it is, in that sense, a measure of
the effect of alkalization upon phytoplankton production.

(It should be

pointed out that it would not be necessary to know the carbon dioxide
production rate in order to calculate the figure in section D* for it
would cancel out in the final subtraction*)
In making the above calculations several assumptions were necess­
ary; they were that (1) the carbon dioxide production observed for the
10-day period (July 15 to July 25) was not higher, due to a more alka­
line reaction and increased decomposition, than for a similar period
of time without alkalization;

(2) there occurred no loss of carbon

dioxide to the atmosphere during the 10-day period for which carbon
dioxide production was calculated, nor during the 8-day period of the
plankton bloom (which is probably a valid assumption since normal car­
bonates were available during both periods for combination with car­
bon dioxide);

(3) the increased standing cron of phytoplankton did not

raise the rate of carbon dioxide production because of the algae’s
respiration, and that (4) the period of plankton production and intense
carbon dioxide utilization was not less than eight days.
It is interesting to note that if any of the above assumptions
were, in faot, invalid, the figure calculated in section D would either
be greater or remain the same; the figure calculated, then, is a mini—

76
vo-Im® •

Furthermore, it would net be expected that "the phytoplankton

would extract the carbon dioxide from all bicarbonate present; in the
eight—day period under consideration, 65% of the available carbon dioxide
was utilized*

If, in section D of the table, 65% of the carbon dioxide

available under conditions of no lime application were subtracted from
the carbon dioxide utilized, the figure in section D would be even larger
(calculated under these conditions, to be 9*4 ppm* COg)*
It can be concluded, therefore, that a causal relationship did
exist between the level of bicarbonate alkalinity and phytoplankton
production during the eight-day period of the plankton bloom*
During 1955, increases in standing crop of phytoplankton were
observed in the treated lakes, particularly in Starvation Lake.

Since,

during the

time of these high standing crops, no reduction in either

half-bound

carbon dioxide or free carbon dioxide occurred, it is not

possible to infer that bicarbonate alkalinity exerted direct effects
upon plankton production by making available more carbon dioxide*

If

the plankton responses were the effect of alkalization (and comparison
with pre-alkalization and control data indicate this to be true), then
it must be concluded that indirect effects (such as greater nutrient
availability) must have been responsible*
The effects of alkalinity upon phytoplankton production may be
summarized in the following hypotheses which received support from the
results of

the present study.

(1) In soft-water lakes the bicarbonate

alkalinity

(or "reservoir" ofimmediately available carbon

dioxide)

limits the rate of carbon dioxide utilization and thus the rate of phy­
toplankton production, but doe 3 not limit the level to which the stand­
ing crop of phytoplankton may reach providing the rate of carbon dioxide
utilization is equal to or less than the rate of carbon dioxide pro­
duction*

(2) Due to the acid conditions and meromictic characteristics

77
(in some c&sss) present in bog lakes* the rate of carbon dioxide pro­
duction is probably less than in more alkaline lakes, which "then limits
the rate of carbon dioxide utilization, or production.

(3) Under opti­

mum conditions of alkalinity, the availability of nutrients, such as
phosphorus, is improved through various mechanisms, and the availability
of such nutrients probably limits, not the rate of phytoplankton pro­
duction, but the level of total standing crop.
Finally, it is possible now to summarize the reasons for observed
differences in production in lakes with different conditions of alka­
linity.

(1) Hard-water, or alkalitrophic, lakes appear to suffer poor

productivity because of the adverse effects of excess calcium upon the
availability of nutrients (see Barrett, 1953).

(2) Medium-hard lakes

with optimum conditions of alkalinity provide large reservoirs of immedi­
ately available carbon dioxide in the form of bicarbonates and permit
phytoplankton ’'blooms” ♦

These lakes, if located regionally such as to

receive nutrients from,for example, fertile soils, will be productive,
but if located such that nutrients are not present, optimum alkalinities
alone will not cause high biological production.

(3) Soft-water lakes,

such as bog lakes, do not permit plankton "blooms" because of a lack of
high concentrations of immediately available carbon dioxide in bicar­
bonates and also limit total production because of the poor availability
of nutrients, whether favorably located in fertile regions or not.
Release of phosphorus from lake soils
The release of phosphorus upon alkalinization may be accomplished
principally by two mechanisms.

The first is by mineralization of organic

phosphorus through decomposition.

Strongly acid conditions, such as

are found in bog lake waters and muds, have inhibitory effects upon
the micro-organisms responsible for decomposition of organic matter and

78
the subsequent mineralization of nutrients*

It is well established that

the addition of liming materials} where such acid conditions exist in
soils, improves conditions for decomposition micro-organisms by increas­
ing the exchangeable calcium and raising the pH and, therefore, hastens
decomposition (see Lyon et alt, 1953, p. 139, and Waksman, 1938, p. 349).
Secondly, phosphate ions may be released through anion exchange;
in the case of the addition of lime, an anion exchange between phosphate
and the introduced hydroxide radicals is probably the exchange involved.
Barrett (1952) in a study of adsorption and regeneration of introduced
phosphorus in lakes, hypothesized that the capacity of lake sediments
for holding exchangeable phosphate is an inverse function of the ratio
of marl to organic matter in the sediments.

He reported that in his

study lakes where such ratios were low, much of the introduced phosphorus
was adsorbed by the sediments and held in exchangeable form.

The above

ratio, for a bog lake, would be extremely low, indicating a high capacity
for the adsorption of phosphate, and it could logically be hypothesized
that increasing the alkalinity in bog lakes would result in a release
of adsorbed phosphate.
Anion exchange involving phosphorus in soils may concern phosphate
adsorbed to clay minerals, organic colloids, and hydrated aluminum and
iron oxides.

It is not likely that clay minerals will be found in a bog

lake, so the sources of exchangeable phosphate are most likely to be
colloidal hydroxides and organic matter.

Colloidal iron hydroxides

are known to exist in lakes (see Ruttner, 1953, p. 74, and Einsele,
1938) and may thus act as a source of adsorbed phosphate; Ohle (1937)
observed an increase in dissolved phosphate after adding a solution of
calcium bicarbonate to an iron-hydroxide gel, and attributed this in­
crease to a decrease of adsorptive capacity due to higher pH values.
In a general discussion of anion exchange in soils, Wiklander

(1955) describes anion exchange mechanisms in "the cases of* clay minerals
and hydrous oxides, and says, "Humic acids "take part in anion exchange,
as evidenced by their power of releasing adsorbed phosphate, but the
exact mechanism is not known#n
In the instances cited in this report where a release of phosphorus
was observed or thought to have occurred (laboratory bottle experimoits
and field experiments on Starvation and Timijon lakes), there was no
evidence to suggest which of the above two mechanisms were in operation#
The bottle experiments definitely showed phosphorus releases, and
results in both Starvation and Timijon lakes indicate releases occurring
from the lake muds*

The most striking increase of total phosphorus in

the field experiments was observed in the bottom waters of Timijon Lake*
where concentrations increased from a mean of 23 ppb* in 1954 to a mean
of 156 ppb* in 1955#

When this increase is considered in the light of

that increase observed in the control lake, Juanita, where total phos­
phorus increased from a mean of 64 ppb* in 1954 to a mean of 80 ppb*
in 1955, also in the bottom waters, some doubt may be cast upon the con­
clusion that the increase in Timijon Lake was due to phosphorus release
from the mud as a result of alkalinization*
tion might be posed:

In other words, the ques­

If an increase occurred in Juanita Lake, the con­

trol, due to natural causes, why could not the increase in Timijon Lake
also be due to natural causes and not associated with lime application
at all?
A graphic comparison of the increases in both lakes is shown in
Figure 28*

First of all, the difference in the two years for Juanita

Lake is statistically significant at the 5% level only and may possibly
be a result of sampling variation alone, while the difference for Timijon Lake is significant far above the 1% level*

Secondly, the nature

of the increase in Juanita Lake between 1954 and 1955, as seen in Figure

80

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•H AJ

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81
is what might he expected, in a meromictic lake under conditions of*
naturally accumulating hypolimnial phosphorus, while the increase in
Timijon Lake was precipitous and of groat magnitude—-an increase which*
if due to naturally—occurring, raeromixis—induced accumulation, would
have happened long ago*
The possibility of the increase in Timijon Lake being due to the
formation of a meromictic layer (monimolimnion) by the density of the
introduced lime solution was discussed in an earlier section, where
evidence was pointed out leading to the belief that the meromictic
character of Timijon Lake was naturally-occurring*
It is tentatively concluded, therefore, from the results of both
the laboratory and field experiments, that phosphorus "would be released
from the lake muds wnen original phosphorus concentrations are low, but
not when phosphorus concentrations are originally high* the postulate
that phosphorus would be released from lake mud upon alkalinization is
supported•
From the practical standpoint, the release of phosphorus due to
increased decomposition would have greater importance than that due to
anionic exchange.

In the former mechanism, the source of phosphorus to

be mineralized may be almost inexhaustable, while in the latter case,
the supply of exchangeable phosphate ions may soon become exhausted, a
phenomenon which has been observed in the use of lime to release nu­
trients in European carp ponds (Neess, 1949)*
The value of applying lime to increase phosphorus concentrations
in hypolimnial waters, such as occurred in Timijon Lake, when such waters
appear never to enter into circulation with upper waters, may be ques­
tioned when such methods are applied in lake management*

The bottom waters

of Starvation Lake (presumably meromictic) held original high concen­
trations of phosphorus; yet biological productivity in this lake (as

82
measured by standing crop of plankton) was no greater than in other bog
lakes lacking such high concentration of hypolimnial phosphorus.

Such

phosphorus release can probably best be utilized in lakes which stratify
yet still are holomictic, or completely circulating; in these lakes,
that phosphorus released in the hypolimnion under deoxygenated conditions
during the summer stratification period would be transported to the upper
trophogenic zone during fall and spring circulation periods*
Color relations in bog waters
The postulate made by Hasler et al* (1951) that color would be
precipitated by lime treatment, is only partly supported by the results
of this study*

A partial flocculation apparently did occur in both

Starvation and Timijon lakes upon alkalization, but this only resulted
in an increase of color and reduction of light penetration*

As was

pointed out previously, the brown strands obvious in unconcentrated
water samples in Starvation Lake were probably the result of partial
flocculation and the cause of the increased color*

The color, at the

time of the increase, was greater at the top of thermocline, than in
the central epilimnion, indicating a settling of the brown strands to
the level of greatest density change, the thermocline.

perhaps if

allcalization had been carried out during times of homothermous condi­
tions in the early spring it would have been possible for the flocculates
of organic matter to settle down completely to the bottom, clearing the
color*
One phase of the color problem, interesting both fundamentally
and from the standpoint of the development of management techniques
for controlling or eliminating color, is the determination of the compo­
sition and source of the color*

In the lakes of the present study,

the color was determined to be of organic nature (as opposed to inor-

83
gQ-nic colloidal iron hydroxides) and composed of particulate matter too
small to he retained hy a fine, quantitative filter paper (Whatman No*
42) yet large enough to he retained hy Seitz bacterial filter discs.
Gorham (1931) was also able to remove color with an ultrafilter, and
concluded that hog lake reactions were due to colloidal organic matter*
In discussing the source of color it is desirable to include also
the subject of acidity, for the two appear to he closely related in
their presence together and may, in fact, have a common source, for the
organic colloids causing color may act as weak acids (humic acids).
Jewell and Brown (1929), in studies of both acid and alkaline hog lakes
in the northern part of Michigan1s lower peninsula, concluded that the
acidity was due to seepage water flowing through the surrounding acid
hog mat, carrying with it organic acids; and that the alkalinity of a
basic hog lake was due to alkaline water entering the lake basin without
passing through the acid-producing part of the hog margin*

Gorham (1931),

in studying the same lakes later, took exception to JeweDl and Brown1s
hypothesis and, on the basis of comparisons among the conditions of the
lake waters, the lake sediments, and material of the bog margins, believed
the source of reaction of the water in both acid and alkaline hog lakes
to be the material at the lake bottom, rather than the margins*

He

believed the colloidal organic matter of the lake water to he derived
from the bottom sediments*
In a discussion of bottom sediments, Ruttner (1953) states the humic
matter leached from the vegetation layer of the soil produces color in
lake waters as a colloidal solution and, upon precipitation, forms dy,
a very finely divided, gelatinous component of the bottom sediments*
The humic matter is thus termed allochthonous by Ruttner*
Welch (1936a) agreed with Gorham on the source of the organic col­
loidal matter*

In discussing a bog lake with highly alkaline waters,

84
he reported the reaction (alkaline, up to pH 8.0) of subsurface waters
observed in near-by springs and other lakes, and also in surface drain­
age streams around the lake in question, as not approaching the basic
reactions of the lake water.

He concludes ,!that an additional contri­

bution of basic—reaction—forming materials must be added after the water
t........................................................................... ....... mm — - — enters the lake basin” (italics Welch*s), and suggests that the basic
bottom sediments are the result of preexisting vegetation (different
from the present acid mat) that produced a basic sort of peat.

Welch

quotes some of Gorham* s chemical determinations where methyl—orange
alkalinities are given as being mostly above 100 ppm., indicating rather
high concentrations of dissolved calcium.

pH values are quoted as being

mostly between 8 and 9, and free carbon dioxide concentrations as be­
tween 2 and 4 ppm.
In the present study, where the lakes appear to be meromictic,
it is difficult to conclude that the organic colloidal material of
the upper waters is derived from the bottom sediment, because the lack
of circulation precludes this source.

Two of the bog lakes studied,

one treated (Timijon) and one untreated (Irwin), underwent a partial
clearing of color from the epilimnial waters which, in the case of Timijon Lake, did not appear to be the result of alkalization (see Figure 18),
and which in the case of Irwin occurred under known natural conditions
(see Figure 23).

No quantitative atmospheric precipitation data were

collected, but the general observations of less rainfall and decreasing
lake levels in the summer of 1955 were easily made.

Another lake in

the same area which was also under study at the same time, and which
had recived lime applications in 1952 and 1953, also was observed to
decrease in color concentration in 1955 (Ball and Waters, MS)»

It would

appear that the lack of run-off water carrying colored organic matter
permitted the decrease in color concentration through natural causes,

85
such as decomposition or settling*

The evidence in the present study

supports the hypothesis of Jewell and Brown (1929) that the source of
the organic matter is in the margin, and does not support the conclusion
reached by Welch and Gorham*
Whether the acid reaction of bog lakes is due to the colored col­
loidal organic matter is still in question#

On almost all sampling

dates for the colored lakes (before alkalization) in the present study
a greater concentration of color occurred in the bottom waters, which
was accompanied by a rise in pH*

Also, the clear-water bog lakes with

low concentrations of color, contained waters with a reaction as acid as
that observed in the colored lakes#

Cowles and Brambel (1934) compared

tests for acidity by titration and for total carbon dioxide by the
manometric Van Slyke method and concluded that about two-thirds of the
acidity was due to carbon dioxide and one-third to colloidal organic
matter, with the test for inorganic acids negative#

Obviously, it is

not sufficient merely to state that the acidity (low pH) observed in bog
waters is due to organic colloidal matter*

The soft waters of most

bog lakes are so slightly buffered that probably any acid, even in
small amounts, will have a great effect upon the pH*
As was pointed out previously, the glacial drift in the area con­
taining the lakes in the present study, being underlain by limestone,
is probably highly calcareous, as evidenced by the presence of limestone
in a near-by gravel pit and the alkaline waters of sj>rings and springfed lakes in the area.

The low content of calcium in these bog lake

waters, then, indicates the presence of an efficient seal against
ground water, probably caused by organic matter in the lake basin*

An

effective seal against alkaline ground water was described in a colored,
soft—water lake, presumably located in calcareous glacial drift, by
Hooper (1954) who reported seepage from an adjacent acid sphagnum bog

86
into the lake*
In the lakes of the present study it is concluded that the source
of color is in run-off water from the marginal areas of the lake (either
the bog mat or uplands in the drainage basin) on the basis of (1) the
decrease of color occurring with decreased atmospheric precipitation,
(2) the impossibility of the source being the bottom sediments in a
meromictic lake, and (3) the apparent seal of the lake basin against
ground water.

Comparison of colored and clear-water bog lakes
In the previous sections several pertinent differences between
the clear-water and colored bog lakes have been pointed out.

These

differences will be summarized in this section, where the data from
the colored lakes, used in the comparison, will be limited to those
from the control, Juanita Lake, and those collected before alkaliza­
tion from the treated lakes, Timijon and Starvation,
Chemical conditions
Conditions concerning dissolved gases were markedly different
in the two types of lakes.

Grant*s Lake did not stratify, so it 'would

be expected that oxygen and carbon dioxide concentrations would be simi­
lar along a vertical series,

Irwin Lake stratified in a manner only

slightly different from the colored lakes, in that the thermocline was
usually about four feet deeper.

Despite the stratification, dissolved

oxygen was found in concentrations suitable for fish well below the
thermocline and, upon several occasions, in the bottom waters.

The

presence of oxygen below the thermocline indicates light penetration
to this depth and the production of oxygen due to photosynthesis.
Hasler's et al. (1951) postulate that oxygen conditions would be im­
proved in the hypolimnion if bog color would be removed is here given

8Y

indirect support.
Generally, chemical conditions relating to types and concentrations
of* electrolytes (pH, alkalinity, total hardness, and conductivity) were
approximately the same in the clear lakes as in the colored lakes*

Total

phosphorus appeared to be slightly higher in two of the colored lakes,
namely Juanita and Starvation, than in the clear lakes; this may be due,
in part, to the higher amounts of suspended organic matter in the colored
lakes.
Color was, of course, much lov/er in the clear lakes and light pene­
tration deeper*

Grant’s Lake was by far the clearest of all studied,

the color usually being about 10 or 15, while Irwin Lake had a color of
40 which was reduced to 15 by the end of the summer of 1955.

No chemical

differences were observed which suggested the reason for less color in
the clear lakes*

In an earlier section concerning color, it was con­

cluded that the source of color was allochthonous organic material de­
rived from the marginal mat or uplands in the drainage basin; it was
observed that in the case of the clear lakes the ratio of bog mat to
water area was much smaller than in the colored lakes*

This suggests

that a correlation between the above ratio and the color of the water,
and possibly even a causal relationship, may exist, out the data are
too few here to attempt such an analysis*
Biological conditions
Qualitatively, there appeared no significant differences among
the plankton of the five lakes, with the possible exception of Grant’s
Lake, where a fewer number of species of phytoplankters were found;
the species found in Grant’s Lake, however, were also found in the other
lakes, both colored and clear*
Quantitatively, also, the standing crop of net phytoplankton ap-

88

peared to be about the same in the clear lakes as in the colored, which
was low*

Standing crop of* zooplankton, however, appeared to be higher

in the clear-water lakes; it is interesting to note that the standing
crops observed in the two treated,

colored lakes after alkali 2ation

were about the same as in the two clear lakes.
Since the fish populations in both clear-water lakes had been
altered artificially, no discussion of the differences in fish popula­
tions will be attempted except to remark on one point.

The possibility of

winter-kill in a colored bog lake is increased because of the large amount
of suspended organic matter, and in the case of meromictic bog lakes, the
deeper waters would not become charged with oxygen at the fall over-turn
(see Greenbank, 1945, for discussion of winter-kill) •

The possibility

of winter-kill may explain the absence of fish in Starvation Lake, which
is highly colored and apparently meromictic.

Winter-kill would not be

as likely in a clear-water bog lake which would jr obably have oxygen
in the deeper waters at the beginning of the winter period and less
suspended organic matter.

Johnson and Hasler (1954) reported winter­

kill in a bog lake which had not been treated with lime while a similar,
near-by, lime-treated lake which had undergone a clearing of color did
not winter-kill, and suggested that lime treatment may prevent winter­
kill in marginal cases by clearing of color.

Such a clearing in the two

treated lakes in the present study did not occur, and an evaluation of
alkalization on this basis, of course, cannot be made.
SUMMARY
To investigate the observed correlation between alkalinity and
productivity of lake waters and to test the use of lime in bog lakes
to increase biological productivity, lime application was made to two
acid, colored bog lakes in the Hiawatha National Forest in the upper
peninsula of Michigan, incorporating a third, untreated, colored bog

89
lake into the sampling program for control purposes.

Two clear-water

bog lakes were also studied for purposes of comparison with the colored
lakes*

Lime application was made twice to each of the two colored bog

lakes, once in the summer under thermally stratified conditions, and once
in the autumn under nearly homo the rmous conditions, by pumping a limewater mixture into the surface waters*

Total rate of lime application

was 100 pounds per acre-foot in each treated lake.
The summer lime applications resulted in raising the pH, and in­
creasing alkalinity, total hardness and conductivity in the epilimnion
only*

The autumn application resulted in similar effects in the lower

hypolimnion, due to a fall of the lime solution to the bottom because of
greater density, which solution did not later circulate with upper layers*
Lime application did not decrease the organic colloidal color, but
produced temporary increases of color immediately following application.
Color decreased the year following lime application in one treated lake
and in one untreated lake, apparently the result of a lack of rainfall
and run-off water carrying colored organic colloids into the lake.
Total phosphorus was greatly increased in the bottom waters of one
treated lake which had had low original total phosphorus content but
did not increase significantly in the bottom waters of the other treated
lake where total phosphorus concentrations were originally high*

Total

phosphorus also apparently increased temporarily in the epilimnion of
one treated lake following the summer application*
Estimations of adsorbed calcium in the bottom muds showed increases
following the autumn application, indicating losses of dissolved calcium
due to cation exchange.
A bloom of nannop lank ton (Microcystis aeruginosa) was observed in
one of the treated lakes after the summer lime application which, it is
shown, because of high bicarbonate content, utilized carbon dioxide at

90
a rate which would not have been possible without lime application*

In

both treated lakes the following summer, significant increases in the
standing crop of net phytoplankton were observed (principally due to
Dinobryon sertularia)*

Increases in standing crop of zooplankton also

occurred in the treated lakes*
Chemical and biological conditions remained relatively constant
in the untreated control lake, and the two species of algae responsible
for the blooms in the treated lakes did not occur at all*
In comparing the clear-water lakes with the colored bog lakes the
principal chemical differences lay in the content of dissolved gases*
One stratified clear-water lake contained dissolved oxygen sufficient
for fish well below the thermocline*

Chemical conditions relating to

dissolved electrolytes were similar to those in the colored lakes before
lime application*
lakes*

Total phosphorus appeared to be greater in the colored

Color, of course, was less and light penetration deeper*

Bio­

logically, .the clear-water lakes appeared to have approximately the
same standing crops of net phytoplankton, but larger standing crops
of zooplankton*

91
ACKNOWLEDGEMENTS
This study was conducted under the direction of Dr. Robert C.
Ball, whose counsel and assistance are gratefully acknowledged.
Appreciation is expressed to the Cutler-Magner Company, of Duluth,
Minnesota, who furnished the hydrated lime used in the experiments,
and to Mr. Robert B* Chapoton who designed and assisted in the con­
struction of the raft-and-puinp system used in lime application.

The

program was financed by the Michigan State University Agricultural
Experiment station, by the Institute for Fisheries Research, Michigan
Department of Conservation, and by fellowships received by the author
from the National Science Foundation*

92

APPENDIX

93
Table 10* Dissolved oxygen, carbon dioxide, and temperature at
all four levels, Starvation Lake**
... ■—

.p. , .iw,, |

Oxygen
ppm
Date
1

8-21-53 5.5
9- 6-53 4.3
5-21-54 8.3
6-28-54 6 . 8
7- 8-54 8.4
7-14-54 5.5
7-15-54 4.4
7-17-54 3.3
7-20-54 3.4
7-23-54 3.0
<*7-25-54 2 . 8
8 - 2-54 10.5
8 - 8-54
7.3
8-18-54 6 . 8
8-28-54 6 . 8
9- 6-54 7.0
10-18-54 5.5
10-21-54 5.7
5-11-55 8.4
6-18-55 7.1
6-28-55 7.4
„7- 8-55 5.5
'7-19-55 5.7
7-29-55 5.7
8 - 9-55
5.7
8-18-55 5.7
8-27-55 3.8

2

Level
3

4.8
1.4

0.0
0.0

8.6

0.7

5.0
2.4
2.3
2.3
1.7
4.0
4.7
6.5
2.3
5.3
3.3
8.7
3.0
1.3
1.5
3.4
9.7
7.0
3.0

0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

2.0

2.5
3.4
4.2
1.3

Carbon dioxide
ppm
4

1

o .c

3
4
3
4

0.0
0.0
0.0

o .c
o .c
0.0

o .c
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
2.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

2
0
0
0
0
0
0
0
0
2
2
1
2

3
2
2
1
2
2

4
3
3
3

Level
2
3 4
4
8

4
8
6
0
0
0
0
0
2
0
1
2
1

3
4
3
2
1
2

5
7
6

5
4
6

7
4
7
8

7
6
6

5
5
6

4
4
5
5
6
6

7
5
3
3
4
5
7
7
7
6
6

Temperature
°P.
Level

18
12
20
20

15
13
15
15
8
12

17
15
15
15
14
12
12
0
0
2
0

3
7
3
8

7
5

1

2

3

4

71
70
60
72
69
72
71
73
71
70
73
72
70
67

69
65
58
69
63
67

44
45
43
48
48
46
46
47
45
46
46
47
45
45
45
45
43
42
43
45
46
45
46
45
45
47
45

44
44
43
46
45
45
45
46
44
45
46
44
44
44
44
43
41
44
43
44
46
45
46
44
45
46
45

68

63
49
48
56
72
70
77
77
75
72
79
72

68
68

65
66
66

69
68

64
65
60
47
46
50
64
68

71
74
73
69
75
70

Air Surface
76
66

69
72
71
67
67
79
60
70
77
69
72
69
67
65
46
57
65
75
74
74
81
78
69
91
72

71
71
58
72
70
72
71
75
71
70
75
72
70
68
68

63
49
50
56
72
71
78
81
75
72
84
71

*Description of sampling levels, as given in text and used in tables
in the appendix:
Level
Level
Level
Level

1,
2,
3,
4,

central epilimnion
top of thermocline
central hypolimnion
bottom ■waters

94
Table 11. pH and alkalinity at all four levels.
Starvation Lake.
Methyl orange
Alkalinity
ppm CaCOg

PH
Date
.1

| 8-21-53
j 9- 6-53
J -5-21-54
6-28-54
; 7- 8-54
I 7-14-54
j 7-15-54
* 7-17-54
{ 7-20-54
5 - 7-23-54
7-25-54
8 - 2-54
8 - 8-54
8-18-54
8-28-54
9- 6-54
10-18-54
10-21-54
5-11-55
6-18-55
6-28-55
? 7- 8-55
7-19-55
7-29-55
8 - 9-55
8-18-55
8-27-55

Level
2
3

5.6
5.5
5.3
5.7
5.5

5.5
5.4
5.1
5.7
5.0

10.8
10.6
10.1

10.6

9.7
9.4
8.5
9.1

9.5
9.4

9.9
10.1

6.6

8.7

5.5
5.4
5.3
5.7
5.3
5.6
5.6
5.7
5.5
5.9
5.4
5.4
5.5
5.0

5.5
5.4
5.3
5.9
5.6
5.6
5.7
5.6
5.5
5.8
5.3
5.2
5.5
5.1

—

-

5.4
5.7

5.4
5.9

6.0
6.6

11.0

8.1

8.0

7.0
7.1
7.0
6.7
6.5
7.0
7.1
7.2
7.0
7.0
6.9
6.9

6.5
7.0

6.8

6 .6

6.1

6.7

6.3

6.5

6.2
6.2
6.1

6.7
7.5
6.9
6.7
6.2

6.4
6.5

4

6.5
6.4
6.7
6.3
6.4
6.6

9.4
7.8
8.1

7.5
7.3
7.7
7.2
7.3
7.5

Level
2
3

1

5
4
4
4
5
22

29
39
41
46
52
42
52
45
41
36
23
23
23
21
21
20
21
20

19
20
20

4
4
4
5
5
19
27
22

Phenolphthalein
Alkalinity
ppm CaCOg

4

1

Level
2
3

12

0
0
0
0
0

0
0
0
0
0

31
31
23

9

46
17 42
16 32
1 1 18

7
5
4
5
5
8
8
10

13
10
12
12
12

24
33
25
46
46
19

6

12

7
7
7
5

13
13 14
16 1

20

6

29
16

10

20

23
22
22
21
21
21
21
22
21

7
16
25
28
27
26
29
27
30
28
29

12

17
13
17
33
81
132
126
133
118
126
134
124
127

12
8

10
6
0

0
0
0
0
0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

7

4

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

0
0

95
35

0
0
0
0
0
0
0
0

0
0
0
0
0
2
0
0

1
f

j

95
Table 12. Total hardness, conductivity, color, and light
penetration at all four levels, Starvation Lake.

Total
Hardness
ppm CaCC>3
Date
1

8-21-53
9- 6-53
^ 5-21-54
6-28-54
v _7- 8-54
7-14-54
7-15-54
7-17-54
7-20-54
7-23-54
/v 7-25-54
8 - 2-54
8 - 8-54
8-18-54
8-28-54
9- 6-54
10-18-54
'"10-21-54
5-11-55
6-18-55
6-28-55
7- 8-55
' 7-19-55
7-29-55
8 - 9-55
8-18-55
8-27-55

5
5
4
4
4
67

Level
2
3

65
63
59
56
54
51
45
41
38
24
25

4
5
4
4
5
42
56
43
29
36
25
48
46
17
19
24
15
16

22

22

68

19
18
18
19
18
17
17
17

19
20

19
20

19
20

17
18

Conductivity
mho X 10"®

4

1

Level
2
3

6

7

4
5
5
5

8

6
6

6
6

7
7

5

5

6

6
6

6

5
5
5
6

5
8

5
5
5
5
8

15
24
24
25
25
25
25
25
25
25

6
6
8
8
6
8
10
10
8
8
12
8
11

118
108
112

113
117
110

115
118
110

118

5
89
83
63
51
54
55
47
44
39
37
32
25
27

47
65
41
25
33
25
42
39
14
16
19
18
18

21
22
21

22
22
22

18

19
23

22
20

19
18
19

21
21

18
20

8
6
6
6
6
8
8
8

7
8
8

9
7
6

7
7
10

17
23
27
28
23
30
27
26
26
27

Light
Penetration
3 ecchi,feet

Color

4

1

2

80 80
80 90
10
80 80
12
85 90
14 90 90
1 2 130 2 0 0
15 130 140
16 125 140
1 1 1 1 0 150
15 1 0 0 130
16 1 1 0 130
16 90 150
14 85 1 0 0
15 80 1 1 0
19 80 1 0 0
14 75 90
19 80 90
147 80 90
91 80 90
130 85 90
85 85
122
118 80 80
139 85 90
80 80
122
125 85 90
119 80 80
125 75 80
13
14

Level
3
80
70
90
110
100
100
100
100

90
90
100
100
110
110
100
100
100
100
100

115
130
150
140
140
150
150
170

4
120
120
120
110

7.5
7.5
7.5
7.0

130

6.0

110
110

5.0'
5.0
5.5
6.5
6.5
6.5
3.0

130
95
110
110

130
130
130
120
120
120
120
120
120

180
170
160
180
160
160
160

6.0

8.5
7.5
8.0
8.0

7 .0
8.0

7.5
8.0
8.0

7.0
6.5
7.5
10.0

9.0

Table 13. Total phosphorus at all four levels and
adsorbed calcium in mud, Starvation Lake*
Total
phosphorus
ppb p
Date
1

8-19-53
9- 6-53
5-21-54
6-28-54
7- 8-54
7-14-54
7-25-54
8 - 8-54
8-18-54
8-28-54
9- 6-54
5-11-55
6-18-55
6-28-55
7- 8-55
7-19-55
7-29-55
8 - 9-55
8-18-55
8-27-55

15
20

9
40
68

24
15
22

23
27
23

Level
2
3

tm
-

Sample
4

25
35
31
29

157
193
226
162

22

201

20

54
34
47
30
19

233
245
280
229
290
226
271
214
249

11

13
13
39
57
15
55
50
45
52

66

68

10

15
13

44
13
19
15

24
76
34
42
34
33
30

12
12

Adsorbed
Calcium
% Ca

5

8

10

14

211

264
249
223

1

0*26
0*24
0*26
0*22
0*22

0*24
0*36
0.26
0*26
0*28
0*22

0*40
0.38
0.32
0.32
0.40
0*42
0.30
0.40
—

2

0.28
0.26
0.24
0.24
0.30
0.24
0.28
0.22

0.24
0.30
0.24
0.40
0.36
0.34
0.42
0.34
0.40
0.32
0.52
—

1

97
Table 14* Dissolved oxygen, carbon dioxide, and temperature at
all four levels, Timijon Lake#
Oxygen
ppm
Date
1

8-16-53
8-31-53
5-19-54
VB 6-27-54
Z - 7-54
7-18-54
7-23-54
7-27-54
c,8 - 2-54
8-12-54
8-15-54
8-24-54
9- 3-54
10-16-54
10-19-54
5- 9-55
6-15-55
6-25-55
*7— 6-55
7-16-55
7-26-55
8 - 5-55
8-15-55
8-25-55

2

Level
3

3.5
6.5 1.4
9.5 10.5
7.3 7.6
7.0 8.0
7.5 7.4
6.5 7.5
6.6

6.0
5.9
7.8
6.8
6.8

8.0
8.2
8.0
10.0

7.9
7.9
7.4

6.8
6.1

7.2
6.5
7.5
6.7

8.1
2.7
7.0
7.9
7.7
9.1

3.0
1.9
1.5

Level
2 3 4

4

1

0.0

3
3
3
3
3
3

4
4

1
1
1
1
1

2

o.c
o.c
0.0 o.c
1.5 0.0
0.0 o.c
0.0 0.0
0.0 0 . 0
0.0 o.c
0.0 0.0
0.0 0.0
0.0

0.0
0.0
0.0

0.0

1.5

4.7

0.6
0.0
0.0
0.0

0.0
0.0
0.0
0.0

0.0
6.8
7.1 0.0
6.8
6.7 3.7 0.0
6.2 4.5 0.0
5.1 2.0 0.0
6.8

Carbon dioxide
ppm

o.c

0.0
0.0
0.0
o.c
o.c

0
0
1
1

2

5
5
5

3

6

3
3

4
5
4
4
4
4
4
5
5
4

1
2
1
1
0
1
1

0 0
1 2 2
2
1
1
2
2 3
2 2 3
2 2 4
2 4
1
2 4 4
2

3

3
5

5
5

Temperature
°P.
1

6

5
5
8
8
8
6

7
12
11
12

9
7
6
0
0
0
0
0
0

0
0
1

4

72
75
56
69
69
75
72
73
70
68

70
72
66

50
50
53
68
66

80
76
77
79
73
73

Level
2
3
66
66

55
67
64
67
67
68
66
66
66
68

64
49
47
48
64
62
72
70
75
73
72
72

50
47
45
46
50
48
47
47
47
47
48
47
45
44
44

41
46
46
47
46
48
47
47
48

4
45
44
44
46
45
46
45
45
45
45
45
46
44
42
46
42
44
46
45
45
45
46
45
46

Air Surface
66

72
49
61
69
75
73
74
66
68

77
80
63
45
44
44
79
64
83
73
84
80
75
74

72
75
55
70
70
74
74
74
71
68

71
72
66

50
54
54
70
66

85
77
78
79
73
73

30

Table 15. pH and alkalinity at all four levels.
Timijon Lake.
Methyl orange
Alkalinity
ppm CaC03

pH
Date
1
8-16-53
8-31-53
^ 5-19-54
6-27-54
7- 7-54
7-18-54
7-23-54
7-27-54
8- 2-54
8- 6-54
8-12-54
8-15-54
8-17-54
* 8-18-54
8-22-54
8-24-54
8-26-54
8-28-54
8-29-54
9- 3-54
10-16-54
10-19-54
5- 9-55
6-15-55
6-25-55
, 7- 6-55
' 7-16-55
7-26-55
8- 5-55
8-15-55
8-25-55

5.6
5.6
4.9
5.8
5*4
5.3
6.7
7.3
7.0
7.2
7.2
7.7
7.9
8.4
6.5
8.2
8.2
7.6
8.9
8.5
7.4
8.2
7.7
7.4
6.6
7.3
7.3
7.2
7.4
7.1
7.0

Level
2
3
5.4
5.4
5.4
5.8
5.5
5.4
5.3
6.3
6.7
5.8
6.9
7.4
7.5
7.8
6.4
8.4
7.3
7.4
8.7
7.0
7.4
10.1
7.1
7.4
6.7
7.4
7.4
7.2
6.9
6.9
6.8

5.4
5.4
4.9
5.7
5.0
5.1
5.1
4.7
5.2
5.1

4
5.3
5.4
5.1
6.1
5.6
5.6
5.4
4.9
5.3
5.3

-

-

5.6

6.0

-

-

—

—

-

-

-

-

-

-

5.4
6.3
9.7
7.0
6.9
7.1
6.6
6.9
6.5
6.7
6.8
6.9

-

5.5
6.1
11.5
9.9
9.6
9.4
8.9
9.1
8.5
8.6
8.3
7.8

1
4
3
2
4
3
3
12
17
16
21
22
27
25
26
29
31
33
32
31
33
23
28
29
28
27
28
26
28
27
26
26

Level
2
3
4
3
2
3
3
3
7
11
11
8
21
20
22
23
23
21
28
30
28
23
22
31
31
29
29
29
31
29
31
27
27

Phenolphthalein
Alkalinity
ppm CaC03

4

1

—

7
7
3
8
8
9
7
9
10
10
11
10
-

—

-

0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
2
1
0
8
4
0
0
0
0
0
0
0
0
0
0
0

3
3
3
5
4
2
4
3
4
6
5
6
5
-

5
9
35
38
41
39
41
41
42
40
41
40

13
-

12
13
29
48
57
59
64
63
68
68
76
71

Level
2
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
0
0
5
0
0
23
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0
0
0
—

0
0
0
12
0
0
0
0
0
0
0
0
0

4
0
0
0
0
0
0
0
0
0
0
0
0
-

0
0
0
146
42
30
15
19
13
6
6
0
0

99
Table 16, Total hardness, conductivity, color, and 1i ght
penetr ation at all four levels, Timijon Lake,
Total
Hardne s s
ppm CaC 03
Date
1
8-16-53
8-31-53
5-19-54
* 6-27-54
7- 7-54
7-18-54
7-23-54
7-27-54
8- 2-54
8- 5-54
A 8-12-54
8-15-54
8-24-54
9- 3-54
10-16-54
"10-19-54
5- 9-55
6-15-55
6-25-55
7 - 6-55
7-16-55
7-26-55
8- 5-55
8-15-55
8-25-55

4
5
5
4
4
4
12
18
17
21
21
26
31
37
26
29
32
30
27
27
26
26
25
25
26

Level
2
3
5
6
5
4
4
4
5
11
11
7
18
21
20
21
24
51
35
30
30
28
31
29
30
27
29

Conductivity
mho X 10- 0

4

1

5
6
5
6
5
5
5
5
5
5
4
5
6
4
5
8
8
5
6
8
7
5
7
5
6
9
7
5
10 11
48 154
46 86
41 80
40 72
40 70
41 69
42 64
41 67
42 67
42 67

6
7
6
5
5
6
12
18
15
20
21
28
30
30
27
30
25
29
27
30
26
29
29
30
27

Level
2
3
6
7
5
5
5
6
6
11
11

8
18
23
19
21
26
53
26
29
29
31
29
31
33
32
30

Li ght
Penetration
Secchi, feet

Color

4

1

6
6
7
8
6
6
6
7
6
7
7 11
7
9
7 10
7 11
7
9
8 12
8 12
7 13
6 11
10 16
49 286
31 56
40 73
39 65
44 77
39 67
44 71
46 77
49 82
43 74

60
60
55
55
60
60
60
60
60
60
60
50
50
55
55
55
55
55
55
50
50
40
45
45
45

Level
2
3
60
80
60
70
65
60
60
60
60
60
60
60
80
60
60

iio
55
55
55
55
55
50
50
45
50

4

60
80
90
60
60
50
90
60
90
60
60 120
60
90
60 110
60 130
60 110
70 120
60 120
70 120
70 110
70 110
100 " 100
55 160
55 160
55 160
65 120
60 110
60 120
60 160
55 160
70 150

9.0
10.0
8.5
7.0
8.5
8.5

8 .S'
10.0
9.0
8.5
11.5
11.0
10.5
7.5
9.0
6.0
8.0
10.0
8.0
8.0

8.0
10.0
11.0
12.5
11.0

Table 17, Total phosphorus at all four levels, and
adsorbed calcium in mud, Timijon Lake*
Total
Phosphorus
ppb P
Date
1

8-16-53
8-31-53
5-19-54
6-27-54
7- 7-54
7-18-54
7-27-54
8 - 5-54
8-15-54
8-24-54
9- 3-54
5- 9-55
6-15-55
"*6-25-55
7- 6-55
7-16-55
7-26-55
8 — 5-55
8-15-55
8-25-55

Level
2
3
_

—

-

-

-

2

6
12
10
10
11

10

3
5
9
4
7
5
5
3
11

24
12

13
7
10
10
8

38
38
14

11
6
6

11
11

15
7
9

7
14
16

Adsorbed
Calcium
% Ca
Sample
4

1
0.22

0.22

0.24
0.24

14

26
29

0*26
0*30
0*26
0.24
0.24
0.24

10

22

0.22

13

11

26
24
26
183
145
154
140
113

14

111

11

142
199
214

0.28
0.24
0.24
0.42
0.36
0.42
0.36
0.42
0,42
0.33
0.38
-

3
16
11

10

7
6
10

36
15

13
11

-

12
22
21

2

0.22

0.24
0.24
0.22
0.22

0.26
0.24
0.26
0.14
0.41
0.34
0.20

0.28
0.30
0.34
0.38
—

'

101
Table 18* Dissolved oxygen, carbon dioxide, and temperature at
all four levels, Juanita Lake*
Oxygen
ppm
Date
8-19-55
9- 3-53
5-20-54
6-21-54
7- 1-54
7-15-54
7-26-54
8 - 6-54
, 8-16-54
' 8-26-54
9- 4-54
5-10-55
6-17-55
6-27-55
7- 7-55
7-18-55
7-28-55
8 - 8-55
8-18-55

Level
3

1

2

4.8
5.0
8.5

2.0
2.0

6.0

6.5
6 .4
6.4
5.2
7.0
5.8
5.9
8.3
7.0
7.4
5.0
5.8
5.3
4.5
5.9

9.4
7.5
7.5
5.3
7.5
5.8
5.3
4.4
2.5
4.9
5.9
4.0
3.1
4.1
3.0
2.7
4.1

3.4
1.9
3.6
IV*
0.4
0.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

Carbon dioxide
ppm
4

1

0.0
0.0

7
5
4
5
4
3
3
4
3
4
4
3
4
3
4
4
4
4
4

o.c
0.0
0.0
0.0

0.0
0.0
o.c
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

Level
3 4
2
12

9
4
5
5

6
6

7
9
9

10

9
14
9
10

6

8

9

5
7
7
87
4
5
4
7

7

10

6
8

11

7
7
6

7
8

7

6
6

8
8

7

9
9

8

9
9
8
8

9
15
13
15
16
13
18

Temperature
°F.
1

Level
2
3

69
79
57
70
72
73
74
70
70
69
63
53
71
69
80
79
75
72
78

60
65
53
64
65
69
65
67
65
64
61
47
67
62
63
74
73
69
75

45
45
45
47
46
46
48
47
46
45
45
42
46
46
47
46
46
45
48

4
43
44
43
45
45
45
45
46
45
44
43
41
44
44
45
44
45
44
46

Air Surface
69
82
60
68

67
73
83
71
64
69
64
51
76
72
84
79
77
71
83

68

78
56
70
72
72
75
70
71
71
63
53
74
69
83
80
76
73
78

102
: table 19* pH, alkalinity, total phosphorus, total hardness,
conduc tivity, color, and light penetration at all four levels,
Juanit a L a k e •
Methyl orange
Alkalinity
ppm CaC 0 3

pH
Date
8-19-53
9- 3-53
5-20-54
6-21-54
7- 1-54
7-15-54
7-26-54
8 - 6-54
_■8-16-54
JuS
\ 8-26-54
9- 4-54
5-10-55
6-17-55
6-27-55
7- 7-55
7-18-55
7-28-55
8 - 8-55
8-18-55

Level
1

2

5.4
5.4
5.0
5.6
5.0
5.0
4.9
4.8
5.0
4.8
5.2
5.0
5.0
5.1
5.1
5.0
4.9
5.2
5.2

5.2
5.2
5.0
5.4
4.8
4.9
4.6
4.9
4.8
4.6
4.8
4.8
4.8
4.9
5.0
4.9
4.8
4.9
4.9

3

5.2
5.2
5.2
5.2
4.9
5.1
5.2
5.7
4.6
5.2
4.8
5.1
4.4
4.7
5.5
5.6
4.9
5.5
4.7
4.9
4.8
5.2
4.8 . 5.2
5.1
4.8
4.8
5.1
4.8
5.2
4.8
5.2
4.9
5.8
4.8
5.1
5.0
4.9

Total
Hardness
ppm CaC0 3
Date
1

8-19-53
9- 3-53
5-20-54
6-21-54
7- 1-54
7-15-54
7-26-54
8 - 6-54
8-16-54
8-26-54
9- 4-54
5-10-55
6-17-55
6-27-55
7- 7-55
7-18-55
7-28-55
8 - 8-55
8-18-55

6

5
4
5
5
5
4
4
4
4
4
5
4
4
5
6

5
5
5

Level
2
3
6
6

4
5
5
5
4
5
4
4
4
5
4
5
5
5
6

5
5

4

5
6

4
5
5
5
5
5
4
4
4
5
5
5
5
5
7
5
5

1

Level
2
3

3
3
3
3
3
3
4
3
4
4
3

3
3
3
3
3
3
5
4
3
3
3

2

2

2

3
3
3
5
5

3
3
3
5
5
5

3
3
3
4
5
5

6

6

6
6

Total
Phosphorus
ppb P

4

3
3

6
6

2
2
2
2

4
4
4

3
4

5
7
5

1

w

8
8

7
5
7
5
7
7
7
14
39
12
10

6

2
2

6
6

3

4
5
5
5
6

6

7
7
7

7
10
7

Conductivity
mho X io~o

4

1

7
7
4

8

6

6

5
5
5
5

7

6

6
6
6

4
5
5
5
5

7
7

8
8

7

7
8
8

6

7

5

8
8

6

5
5

7
8

Level
3
2

4

1

90
90
90
80
85
85
80
90
90

8
8
8

7
9

6
6

7
7

7

7

8

8
8

8
8

9

9

8

8

8

7
7
7

7
7
8

10

11

11

8

5
11
13
14
15
14
13
17
40
11
17
11
15
11

17
17
15
15
13
13
15
23
26
22
17
20
14
14
15

8

8

8

8
8
8

7
9
^

7
9
9

8
8
8
8
8
8

9

8

8

8

8

9

100

90
90
100
100

90
80
85
90
80

2

Level
3
120

120

90 110 160
1 2 0 160
90 100 170
90 110 150
130 120 180
140 120 140
130 120 150
120 120 140
100 120 150
95 100 125
100 120 170
140 120 200
120 130 170
95 100 140
120 120 150
120 120 150
100 110 140
100

mm

55
61
63
54
73
79
69
64
61
48
101
69
83
85
86

79
87

4

180 130 130
200

4

Light
penetration
Secohi.feet

Color

7v
9^
7

7
7
7
7

Level
2
3

7.0
.5
5.0
7.0
7.0

6

8.0

7.5
7.5
7.0
8.0

6.5
6.5
7.0
7.0
6.5
7.0
6.5
7.0
8.0

103
Table 20. Dissolved oxygen, carbon dioxide, temperature, pH,
alkalinity, total phosphorus, total hardness, conductivity,
color, and light penetration at all four levels,
Irwin Lake*

Oxygen
ppm
Date

Level
2
3

1
8-22-53
9- 1-53
6-26-54
8-13-54
9- 5-54
6-23-55
7-16-55
8-26-55

Carbon dioxide
ppm

7.4
6*5
7.5
8.0
8.5
8.5
8.0
7.6

7.0
6.0
8.5
8.7
7.8
9.2
8.3
8.0

5.8
5.2
7.5
5.5
4.0
4.9
6.5
2.1

4

1

Level
2 3 4

1

Level
2
3

4

3.1
L.2
I5.5
0.0
0.0
0.3
0.0
0.0

3
3
3
2
2
2
2
2

3
3
3
2
3
2
2
2

74
77
71
69
67
67
77
73

70
72
68
62
64
64
73
71

45
47
48
47
47
44
46
46

Level
2
3

1
8-22-53
9- 1-53
6-26-54
8-13-54
9- 5-54
6-23-55
7-16-55
8-26-55

5 .6
5 .5
5 .6
5 .0
4 .8
5 .2
5 .1
5 .1

5.4
5.4
5.4
4.9
4.9
4.9
4.9
5.0

5 .2
5 .3
5 .3
5 .2
4 .9
4 .9
4,»9
4 »8

8-22-53
9- 1-53
6-26-54
8-13-54
9- 5-54
6-23-55
7-16-55
8-26-55

1

Level
2
3

4

1

4
5
5
4
3
4
4
4

4
6
4
4
2
4
4
4

4
6
5
4
5
4
4
5

8

4
6
4
3
2
4
4
4

4
4
4
4
5
3
5
5

50
53
50
50
48
45
54
48

1

Level
2
3

4

1

5.2
5.3
5.3
5.6
5.3
5.2
5.1
5.7

3
3
2
3
3
2
4
2

3
3
3
2
2
2
4
2

3
3
3
2
2
2
3
2

3
4
2
2
5
3
4
6

—
5
0
5
3
6
2

8
7
8

9
6
7
7

Level
2
3
8
8
7
8
7
6
7
8

8
8
8
8
7
6
9
7

1

8
9
8
8
7

40
40
40
40
30
25
20
15

6
8
8

74
77
70
70
67
67
77
73

4

—
2
8
15
10
8
21

6
60
74
44
46
54

2
6
11
5
4

Light
Penetration
Secchi,feet

Level
3
2

4

25
30
40
30
40
30
20
20

25
30
40
50
70
50
50
75

40
40
40
40
40
25
20
20

77
82
76
73
75
62
75
69

Level
2
3

4

Color

4

Air Surface

Total
phosphorus
ppb F

4

Conductivity
mho X i o - b

Total
Hardness
ppm CaCOg
Date

4
4
4
3
4
3
3
4

Methyl orange
Alkalinity
ppm CaC03

pH
Date

Temperature
°F.

12.5
14.0
14.5
13.0
13.0
13.0
13.0
15 .0

1U4

Table 21* Dissolved oxygen, carbon dioxide, temperature, pH,
alkalinity, total phosphorus, total hardness, conductivity,
oolor, and light penetration at all three levels,
Grant»s Lake.

Carbon
dioxide
ppm

Oxygen
ppm
Date
8-24—53
9- 5-53
6-20-54
8-12-54
9— 4-54
6-20-55
7-15-55
8-25-55

1

Level
2

3

1

Level
2 3

7.7
7.4
8.0
7.8
7.8
8.2
7.7
6.9

7.8
7.3
8.0
8.2
7.8
8.5
7.5
6.9

7.8
7.1
9.0
7.8
7.8
8.3
6.3
0.6

2
2
2
1
2
2
2
2

2 2
2 2
3 3
1 2
2 2
2 3
2 5
2 12

73
73
71
69
66
73
77
77

72
73
70
70
67
68
76
76

1

Level
2

3

1

Level
2

3

5.7
5.7
4.8
5.0
5.3
4.5
4.9
4.9

5.6
5.6
4.8
5.3
5.4
4.5
4.9
4.9

5.6
5.9
4.5
5.1
5.4
4.4
4.8
4.9

3
3
1
2
2
2
2
2

3
3
2
3
2
2
2
3

3
4
1
2
2
2
2
4

Total
Hardness
ppm CaC 03
Date
1
8-24-53
9- 5-53
6-20-54
8-12-54
9- 4-54
6-20-55
7-15-55
8-25-55

Level
2
3

1

4
3
4
3
3
3
4
4

Conductivity
mho X 10-6

Level
3
2

1

4
4
4
3
3
3
4
6

8
7
7
7
5
8
7
7

4
4
4
3
3
3
4
5

72
72
58
70
67
60
68
71

Methyl orange
Alkalinity
ppm CaC 03

PH

8-24-53
9- 5-53
6-20-54
8-12-54
9- 4-54
6-20-55
7-15-55
8-25-55

Temperature
°F.

74
63
71
67
62
79
76
79

Surface
73
74
71
70
67
72
77
77

Total
Phosphorus
ppb P
1

-

1
0
3
0
0
0

1

Level
2

7
5
7
7
6
8
7
8

10
10
5
15
20
15
15
10

10
10
10
15
20
15
15
15

Level
2

3

—

—

0
0
7
2
4
0

0
2
6
5
20
63

Light
penetration
Secchi,feet

Color

Level
3
2
7
6
7
7
6
8
7
8

Air

3
30
40
15
15
20
15
20
40

19
16
20
16
13
22
19
13

1UD

Table 22. Vo lurne of plankton, in cubic milimeters
X 10- 3 per liter,
Starvation Lake •

Phytoplankton
Excluding
Peridiniiurn
Date
8-21-53
9- 6-53
5-21-54
6-28-54
7- 8-54
7-14-54
7-25-54
R—
O

R — Oft
RA
O

R—
O

R—0
R^
A
0

R
1R RA
O — iu-

8-28-54
Q
R — oft
CA
y— o

5-11-55
6-18-55
6—28—55
7- 8-55
7-19-55

8- 8-55
8-18-55
8-27-55

Level
1
2
1
2
1
2
1
2
1
2
1
2
1
2
1
2
1
Z.

Sample
1
2

1
4
8
5
9
13
4
0
1
4
0
0
0
0
0
2
0
0
5
1
2
0
1
1
2
1
1
1
16
1
0
2
0
217
1
2
48
1
134
2
73
149
1
2
63
1
455
359
2
1 4653
2 2265
1 1515
2
576
1
0
2
11
10
~"T~
2
6

0
8
11
4
6
12
3
0
1
0
0
0
0
0
0
2
0
0
1
2
1
1
10
28
0
0
212
55
42
62
193
67
504
273
3439
1924
1250
593
1
14
1
8

Mean
1
6
10
5
8
13
4
0
1
2
0
0
0
0
0
2
0
0
3
1
1
1
6
22
0
0
215
52
88
68
171
65
480
316
4046
2095
1382
585
1
13
5
7

Zooplankton
Peridinium
only
Sample
1
2
17
17
423
51
0
0
135
68
68
85
0
338
0
321
0
85
17
17
85
0
0
0
34
0
0
0
85
17
0
0
0
0
17
0
0
0
0
0
0
0
0
0

17
34
473
101
0
0
203
118
152
68
0
169
0
220
0
34
17
135
17
0
17
0
0
0
0
0
34
17
17
0
0
0
0
0
0
0
0
0
0
0
0
0

Mean
17
26
448
76
0
0
169
93
110
76
0
254
0
270
0
59
17
76
51
0
9
0
17
0
0
0
59
17
9
0
0
0
9
0
0
0
0
0
0
0
0
0

Sample
1
2
608
291
330
136
256
86
289
408
205
438
0
81
0
0
11
5
501
137
46
103
706
818
1459
2230
1077
97
2400
581
1127
1585
2343
1395
1287
1983
860
791
1485
900
1095
1445
2016
693

560
803
310
0
423
68
544
319
73
570
0
40
0
45
11
0
137
137
23
171
1680
942
1307
1306
1522
92
1075
499
1639
634
2601
1644
2318
884
583
1219
684
586
556
1206
1063
1033

Mean
584
547
320
68
340
77
416
363
139
504
0
60
0
23
11
3
319
137
34
137
1193
880
1383
1768
1300
95
1738
540
1383
1109
2472
1519
1802
1433
721
1005
1084
743
826
1325
15 39
863

106
Table 23. ¥01111116 of plankton, in cubic milimeters X 10“5 per liter
Timijon Lake.
r

Phytoplankton
Zooplankton

Excluding
Peridinium
Sample
Date

Level

8-16-53
8-31-53
5-19-54
6-27-54
7- 7-54
7-18-54
7-27-54
8- 5-54
8-15-54
8-24-54

1

2

9- 3-54
5- 9-55

6-15-55
6-25-55
7- 6-55
7-16-55
7-26-55
8- 5-55
8-15-55
8-25-55

1
2

Peridinium
only

Mean

1

2

1
6
0
6
52
58
36
31
13
13
5
5
2
1
1
4
2
1
2
0
0
4
0
31
52
47
161
30
523
297
261
246
420
725
222
254
10
5
17
4

0
7
4
5
32
29
29
25
5
16
3
7
1
0
3
4
2
1
0
2
0
10
0
31
55
55
123
38
424
172
339
339
356
469
280
235
5
10
9
26

1
7
2
6
42
43
33
28
9
15
4
6
2
1
2
4
2
1
1
1
0
7
0
31
54
51
142
34
473
235
300
292
388
596
251
244
8
8
13
15

Sample
1
2
51
17
17
0
0
0
0
17
17
0
17
0
34
0
0
17
17
34
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
17
34
68
85
34

51
17
17
0
0
0
0
0
0
0
17
17
17
17
0
0
34
0
0
0
0
0
17
0
0
0
0
0
0
0
0
0
0
0
0
0
51
51
135
68

Mean
51
17
17
0
0
0
0
9
9
0
17
9
25
9
0
9
25
17
0
0
0
0
9
0
0
0
0
0
0
0
0
0
0
0
0
9
42
59
110
51

Sample
1
2
667
56 3
130
589
238
282
234
343
313
392
737
386
52
59
153
209
235
602
357
717
134
804
35
194
936
2116
1397
1055
656
418
778
1327
952
1005
1665
998
2146
1962
2176
853

650
744
185
694
388
288
328
372
327
376
636
209
34
268
69
177
309
766
226
630
161
810
148
180
1334
1318
436
1068
633
632
975
1192
789
878
2448
762
1888
2297
1765
2166

Mean
658
653
158
641
313
285
281
357
320
384
686
298
43
163
111
193
272
684
291
673
147
807
92
187
1135
1717
916
1061
644
525
877
1259
870
942
2057
880
2017
2130
1971
1509

107

T£at a ^ a

l:^e0 f plaaktaa* in

- l - e t ers X xo-3 per l i t e r , |

Phytoplankton

Date

Level

Peridinium
only

Sampl e

Sample

1

9- 3-53
5-20-54

2
1

7
9

10

8

5

7

6
12

6
11

25

25
3
15
5

10

7-15-54

4
7

7-26-54

2

8- 6-54
8-16-54
8-26-54
9- 4-54
5-10-55
6-17-55
6-27-55
7- 7-55
7-18-55
7-28-55
8- 8-55
8-18-55

1
2
1
_2_
1
2
1
2_
1
_ 2_
1
_2_
1
2
1
_2_

\
2

4

3
1
6
10

5
25
4

7- 1-54

1

1

10

6-21-54

Mean
2

4

8-19-53

Zooplankton

Excluding
Peridinium

2
20
6
12

Mean
2

0
0
0
0
0

0
0

0
0

17

9

0
0
0

0
0
0
0
0

0
0
0
0
0

34

17

25

0
0
0

0

0

17

9

0

0

17
17

17
17
0

0

51

25

0

0

6

0
0
0
0
0

17
9
9

34

17

1
10

2
10

0

0

0

372

152

4
7
7

4
5
7

68
0
0

101
0
0

262
85

2

2

2

51

17

34

2
12

3
15

0

0

51

0
68

7
4
14
33

4
18
9
16
9
18

8
10
12

304
34

8

21

46

24
15

30
14
15
4
6

7
3
10

3
2

7

10

31
26
102
101

13
14
17
13
10

4
4

20

15

10
8
22

15
14
7
5

25
15
35
13
26

121

21
111

78

89

0

0
0

202
0

85
34
254
34
169
34

17
279
34
186
17

118

101

110

17
152
152
490
34

9
144

0

0

135
51
490
152

101

490
93

Sample

Mean

1

2

479
76
514
84
460
253
581
231
283
204
218
137
77

339
19
345
123
789

0

116
189
114
218
141
92
449
657
257
58
419
271
421
700
896
1161
605
629
610
390
368
613
1121

872

202

292
145
178
183
228
211

118
69
60
325
121

84
110

90
326
535
216
251
355
402
493
282
1055
721
1021

858
955
413
502
417
1639
983

409
47
429
103
624
227
436
188
231
194
223
174
98
35
88

258
118
151
125
91
388
596
236
154
387
337
457
491
975
941
813
744
783
402
435
515
1380
928

108

Table 85 • Volume of plankton, in cubic milimeters X 10" 3 per liter,
Irwin Lake*

phytoplankton
Excluding
Peridinium
Sample
Date

Level

0“ 6&"wu
Cl
y— 1i-od
Q

6-E6-54
8-13-54
9- 5-54
6—23—55
7-16-55
8-26— 55

1
2
1
2
1
2
1
2
1
2
1
2
1
2
1
2

Peridinium
only

Mean
2

1
6
0

7
7
23

5

0

0

0

1
20

1

0
0
0
0

0
0

0
0

19
1

104

36
15
239

111

2

120

272
349
96
124

14
13
12
10
2
2

70
14
247
170
331
441
82
117

0

0

0
0

0

0
0
0
0

17

17

17

0

0
101

0

17

85
34
17
85
0
0

Sample
1

4

9
3

12

Mean
2

1

10
1
2

254
219
390
532
69

Sample

9
0
0

0

93
25
17
42

0
0

0
0

17
17

1349
959
1322
3108
3143
2026
913
933
1737
403
518
869
665
979
1001

1014

Mean
2

1586
809
2441
2655
3641
2497
881
256
1657
1190
1689
1314
409
645
307
1255

1467
884
1881
2881
3392
2262
897
595
1697
797
1104
1092
537
812
654
1134

109
Table 26. Volume of plankton, in cubic milimeters X
Grant *6 Lake.

Phytoplankton
Sample
Date

L evel

8-24-53
57— 5*00
6-20-54
8-12-54
9- 4-54
6— 20— 55
7-15-55
8— 25—55

1
2

3
18

7
18

5
18

1
2
1
2

0

0
0
6
2
2

0
2

3

2
2
0

2

2
1
2
1

0
2
0

2
1

1
10

2
1
2

4
23
60
87

Sample

Mean
2

i

1

1
0
1
8
10

18
123
68

p er liter.

Zooplankton

1

3
7
4

10*3

7
3
2

3
5
10
21

92
77

678
2269
1454
795
1209
777
1434
1044
2767
2671
715
1046
776
2399
581
1369

Mean
2

693
2174
945
567
882
1048
1038
1335
2208
1576
465
642
805
2142
1041
2142

686
2222

1199
681
1046
913
1236
1190
2488
2124
590
834
791
2271
811
1756

110

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1955
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1947
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1954
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1936
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1938
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1948
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1945
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1951
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1929
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1954
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1938
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1950
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1953
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1949
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1937
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_
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