A STUDY €31“ PRWIARY PRDDUCTEC‘N @N ARTlFltlAL
SUBSTRATES- IN A REFFLE AND 9031. AREA OF THE
RED CEDAR RIVER

1?th kw Hm aw a; M. 8..
MKFCHGGAN STAGE UNWEMR‘Y
Rab“? Raymond Rawsfmm k.
195%

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A STUDY OF PR MARY PRODUCTIOK OI ARIIFICIAL SUBSTRAIES
IN A RIFFLE AHD POOL AREA OF THE RED CEDAR :IVER

By
ROBERT RAYMOND RAWSTRON, JR.

AN ABSTRACT

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIEXCE
Department of Fisheries and Wildlife

1961

.1 A , /"‘
Approved (/2122 JL (M

 

ABSTRACT

Studies of the primary production of both riffle and
pool situations, using artificial substrates in the Red
Cedar River, were carried out in the summer of 1960. Trans-
plants from the pool to the riffle were made. Primary pro-
duction was measured by the accumulation of phytopigments
measured as AA x 103. PhytOpigments were extracted in 95
percent ethanol and "read" on a Klett-Summerson colorimeter.

The riffles were found to be more productive, attain-
ing higher maxima, and showing faster growth rates than the
pool. The pool substrates reached their maximum standing
crops within fifteen day cycles, whereas the riffle sub-
strates did not. Transplants from the pool to the riffle
showed an increased growth rate after the standing crop
from the pool was accumulated for nine days. Current ve-
locities between 1.0 feet per second and 3.0 feet per sec-
ond showed higher growth rates on the artificial substrates
than that of currents above or below these values.
' Community composition differed; Melosira s . being
dominant in the riffle community, while Synedra ping and
Navicula cryptocephala characterized the pool community.

R. R. R.

 

A STUDY OF PRIMARY PRODUCTION OI ARTIFICIAL SUBSTRATES
IN A RIFFLE AND POOL AREA OF TEE RED CEDAR RIVER

13}!
ROBERT RAYMOND RAWSTRON, JR.

A THESIS_

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIEHCE

Department of Fisheries and Wildlife

1961

 

l5! 7/
7/3/91

ACKNOWLEDGMENTS

The writer wishes to express his appreciation to Dr.
Robert C. Bell for the interest and help he gave in both
the research and writing of this thesis. In addition, the
writer's fellow graduate students in the Fisheries and
Wildlife Department, particularly Kenneth Linton and Dar-
rell King deserve thanks for their help in the field and
their constructive criticism.

The research was financed by the Agricultural Experi-

ment Station of Michigan State University.

11

TABLE OF CONTENTS

INTRODUCTION
DESCRIPTION OF STUDY AREA
METHODS AND TECHNIQUES
Periphyton Sampling Procedures
Current Velocity Measurements
Qualitative Methods
PhytOpigment Extraction and Measurement
RESULTS AID DISCUSSION
Comparison of Standing Crops from Pool and Riffle
Results of "Transplanting"
Effects of Different Velocities
SUMMARY
Appendix I
Matched Pairs Test--Three day
Matched Pairs Test--Six day
Appendix II
Current Velocity Data
Appendix III
PhytOpigment Extract Data
LITERATURE CITED

iii

 

FIGURE
1.

2-8.

10-16.

LIST OF FIGURES

Phytopigment Correction Graph

Comparison of attained standing creps
of pool and riffle for each cycle

Grand average of standing crOps of
pool and riffle

Comparison of standing crop changes
between pdol control and "trans-
plants" at each site for each
cycle

iv

PAGE
13

19-25

30

32-38

NTRODUCTION

The measurement of productivity is fundamental to the
science of limnology. Productivity occurs on two levels.
Primary productivity is defined by Odum (1953) as the rate
at which energy is stored by the activities of primary pro-
ducer organisms in the form of organic materials which can
be used as food. Secondary productivity occurs as the con-
sumer organisms utilize the energy produced on the primary
level.

Most studies to date have centered on lentic environ-
ments, and much remains to be learned of the lotic situa-
tions. Odum's (1956) work in studying primary productivity
of flowing waters in the natural artesian springs, using di-
urnal gas curves and community metabolism methods, is not
generally useful in the warm-water streams in Michigan.

The effects of rapid water temperature change, organic pol-
lution and extreme variability in stream flow introduce
other variables for which it is difficult to account, using
Odum's methods.

The introduction of artificial substrates into both
lentic and lotic situations is not new, and Cooke (1956) de-
scribes some of the history. Many substances have been used
as artificial substrates. Keup (1958) used cedar shingles;
others have used glass slides, stones, cinder bricks and
plexiglass plates to collect the organisms that have become
attached to them.

Much terminology has arisen to describe the complex of

l

2

organisms which become attached to underwater objects. News
combe (1950) and Cooke (1956) give definitive discussions of
these terms. Ruttner's term "aufwuchs" seems to be more
nearly correct than the others, referring only to the organ-
isms which become firmly attached, but do not penetrate in-
to the substrate. Grzenda and Brehmer (1960), using the
same general techniques as in this study, use the term "pe-
riphyton" to describe this assemblage of organisms. Either
term appears to be acceptable, but for the purposes of this
study the term "aufwuchs" is preferred.

Recent use of plexiglass plates in studies at Michigan
State University has demonstrated their validity as an arti-
ficial substrate (Peters, 1959). Investigations so far,
however, have taken no cognizance of possible differences in
productivity levels between pool and riffle conditions on
such substrates. Stokes (1960) in his study of an artifi-
cial stream demonstrated slight differences between the two
situations. Ruttner (1953), and Whitford (1960) discuss the
"physiological richness" of current situations and demon-
strate that differences do exist between productivity of
pools and of riffles.

Peters (1959) gives two assumptions that must be made
when using artificial substrates. "(1) The substrates are
not selective for specific organisms and (2) the production
on the artificial substrate is at the same rate as occurs on
a natural substrate." For this study, the second assumption

will be expanded to read, "the production on the artificial

3

substrate in a given environment (pool, riffle) occurs at
the same rate as on a natural substrate in that same envir-
onment."

This study was carried on to determine whether differ-
ences in rate of production occurred between the pool and
riffle conditions on artificial substrates. Primary produc-
tivity was estimated for each situation by the accumulation
of plant material as measured by phytopigment extracts over
a period of time. Comparisons were made while trying to
keep all other variables as nearly the same as possible,
e.g. water temperature, available light, and nutrients, tur-
bidity, and stream depth. Using the same standards for mea-
surement, plexiglass plates were transplanted from the pool
to the riffle to determine what the effects of velocity
would be on the pool-grown communities.

Only limited taxonomic or quantitative work was done
and this to determine whether the community structure of

each community was similar.

rear-u

 

 

DESCRIPTION OF STUDY AREA
The Red Cedar River is a typical southern Michigan

warm-water stream, characterized by slow currents and pools
with occasional rapids. The upper portions of the river
have been dredged to straighten and deepen the channel while
the major tributaries have been dredged for agricultural
purposes. Both treated and untreated sewage enter the river
throughout the watershed. Three artificial impoundments are
located on the main river; in the town of Williamston, at
Ferguson Park in Okemos, and on the Michigan State Univer-
sity campus.

The Red Cedar River begins at Cedar Lake in southeast-
ern Livingston County in Sections 28 and 29. It runs a
northwesterly course for approximately 18 miles, and then
flows westward through Ingham County for 28 miles to its
confluence with the Grand River within the city of Lansing.

The climate, geology, soils and land use practices are
described by Meehan, 1958.

The sampling stations were located in the main stream
0.5 miles below the bridge at Dobie Road in Okemos. This
choice of sites was made to take advantage of a strong rif-
fle immediately adjacent to a long shallow pool and the
presence of a water temperature thermograph. The stream at
the sampling sites is approximately forty feet wide. The
pool bottom is generally of thick sand, but at its down-
stream end gives way to large rock and gravel. A natural

riffle occurs here, but the water flow was too low, so a

a

double rock wing dam was built to increase the flow and
consequently the velocity, in the middle of the river.

Station A was located in the pool 15 feet from the
south bank of the river. The pool depth at this point is
about three feet. Velocities ranged from 0.168 feet per
second to 0.893 feet per second.

The riffle is approximately 35 feet long by ten feet ‘
wide, sweeping into another large pool. The bottom is of
large rock and debris, and has a nearly uniform depth of
three feet, varying with river flow. It lies approximately
sixty feet downstream from the point at which Station A
‘was established. Its velocity during the study ranged from
h.55 feet per second at its upstream end to 1.23 feet per
second at its downstream end. Station B was established
within this riffle.

The stations were chosen to provide similar ecologi-
cal variables, leaving current velocity as the only recog-
nized difference. Since the stations were so close to-
gether, turbidity, nutrients, and water chemistry were as-
sumed to be similar. Both situations had similar shade
cover. Both were covered most of the morning, and exposed
two to three hours each afternoon, and shaded again in late

afternoon and evening.

We If,

 

METHODS AND TECHNIQUES

The measurement of productivity can be carried out by
many methods. At present more emphasis seems to be placed
on a general method involving the measurement of photosyn-
thetic activity rate through the quantitative estimation of
gas production in light and dark bottles orthe addition of
various inorganic radioactive isotOpes into the medium and
measurement of it after fixation into organic form. C1“,
P32 and other radioactive isotOpes which are fixed in organ-
ic matter have been used.

Productivity can be measured by at least two other genp
eral methods. One involves the estimation of plant material
growth and includes such techniques as the counting of cells
within a unit volume or surface area, the isolation of sin-
gle or mixed components characteristic of plants, the iso-
lation of a single plant constituent, chlorophyll and/or
other pigments having associated optical properties. The
other involves measurement of the total weight of organic
matter accumulated over a period of time. Often all or part
of these general methods are combined in a single study.

It has been demonstrated that artificial substrates can
be used to sample productivity. Peters (1959) establishes
the validity of their use in the Red Cedar River. Cooke
(1956) and Newcombe (1950) review the literature concerning
fresh water community types and discuss the role of the auf-
wuchs community. The aufwuchs are those organisms except

macrophytes which attach themselves to a substrate, but do

6

7

not penetrate into it. Other terms are sessile benthos and
periphyton. Benthos is a large category separating organ-
isms on the bottom from those found free-floating or drift-
ing as plankton. Sessile is applied to organisms which be-
come attached to a substrate whether plant or animal. Peri-
phyton is a term, often used synonymously with aufwuchs,
which may include dead or alive plankters or drifting or-
ganisms of plant or animal origin caught in the attached
forms on a substrate. Since this study is concerned only
with attached forms and not the organisms caught up in them,
the term aufwuchs is preferred.

Plexiglass plates were used as the artificial sub—
strates. Each unit presents l.h square decimeters surface
area for attachment. These plates are quite inert chemic-
ally. They were attached to wooden racks, which in turn
were bolted onto steel fence posts, which had been driven
into the river bed. Grzenda and Brehmer (1960) describe
the technique and give pictures of similar apparatus.

During the summer of 1960 Stations A and B were com-
posed of three sets of ten plexiglass plates, each on racks
placed on the previously installed fence posts. Each post,
with its attached rack, was designated a site. These sites
were given the numbers 1, 2 and 3 to show from which rack a
given plate had come. The plates, in turn, were given code
numbers to indicate the position on the rack from which
it had come. All those on the left as faced from behind

were given odd numbers in order of increasing magnitude

are 1 fr"

finn’.

 

8

from the and toward the middle, while those on the right
were given even numbers in the same way. In addition, a
subscript was given to each code number to indicate Station
A or B. Thus, 16A was the third plate from the right at
Site 1, Station A, while 23B was the second plate from the
left at Site 2, Station B. The shingles at Station A were
considered to be matched working inboard from each end.

At zero day, the beginning of a cycle, the 60 plates
were lowered to 0.8 of the depth from the surface. This
depth was chosen to eliminate the effects produced by sharp-
ly reduced current at the bottom (Welch, 1952), while at the
same time allowing for possibly lowered flow. On the third
day at Station A, the whole rack at each site was raised
and the outer plates removed (11A, 12A; 21A, 22A; 31A, 32A).
One of each of the pairs was taken to the laboratory; the
other was kept wet and handled as little as possible and
"transplanted" to the same position at the same site at Sta-
tion B, whose corresponding plate had been removed prior to
that of Station A.

To the Open position at the sites at Station B a clean
plate was placed. This plate was coded XYbla where X and
Y represent the site and position number respectively. They
were placed here to try to determine the comparative growth
rate in the riffle for the transplant period of a given
shingle.

The procedure moving inboard at Stations A and B con-

tinued every third day for fifteen days. In addition, to

the regular removals, the previous transplant and the plate
coded XYbl were removed. Thus each transplant Spent three
days in the current and either 3, 6, 9 or 12 days in the
pool, while all the XYbl's Spent three days in the riffle.
The preceding technique was carried out from July 2, 1960
to October 17, 1960.

To insure that only the aufwuchs were measured and not
drifting phytoplankters either dead or alive, which had set-
tled out, each shingle was thoroughly stream-washed to re-
move the loose materials. The plates were all washed four
times to add some constancy to the maneuver.

The statistical relationships between the pairs assumed
to be matched were carried out. Two separate matched pair
tests were run in the pool; one for six days, and the other
for nine days. The results of these tests are shown in Ap-
pendix I.

During the summer of 1959, a different scheme was used.
The stations were the same, but Station A had only two sites
while Station B had three. Each site was composed of two
shingles and all positions at each site were removed and
brought into the laboratory every eight days. No trans-
plants were made. Direct comparisons were made for each
period. Little use of these data can or will be made in
this paper. During the eight-day interim the operator of
the impoundment at Williamston often reduced the stream flow
considerably, covering and uncovering the shingles for var-

ious periods of time. Either through experience or more

 

 

10

frequency on the stream during the summer of 1960, this con-
dition was evaded completely, and more useful and quantita-
tive information was obtained.

Current velocity was measured with a Gurley bucket-
type current meter, Model 622 and the micro-unit Model
625-F. Three readings were made directly over the rack on
each side of the fence post at each site and averaged for
each site. Four distinct gradations of velocity were noted.
Therefore, four general classifications were set up. The
pool which always registered velocity less than 1.0 feet
per second and greater than 0.16 feet per second was desig-
nated V0. The riffle area velocity at the three sites was
more varied, but in general Site 1 ranged between 3.5 feet
per second and h.k feet per second. Site 2 ranged from
1.77 feet per second to 3.08 feet per second. Site 3 ranged
from 1.23 feet per second to 1.93 feet per second. Three
categories were set up for Station B and are as follows: 1.0
feet per second to 2.0 feet per second, designated as V3, at
Site 3; 2.00 feet per second to 3.00 feet per second, desig-
nated as V2 at Site 2; and 3.00 feet per second to H.55 feet
per second at Site 1, designated as V1. There was some
overlapping in these categories, but in general they repre-
sent the true picture of the velocities present during the
study. Appendix II shows the average currents present at
each site and the dates.

During the summer periods as described above, the arti-

ficial impoundment at Williamston is in operation. water

T"???
_ ‘

 

11

flow, and hence stream velocity, varied greatly with no a-
parent regularity. In addition, meteorological conditions
caused fluctuations. In order to cope with such large
changes over such short periods of time, it was felt that a
random measuring of the current velocities would be more ef-
fective than trYing to measure every day or at regular in-
tervals.* Ten random numbers were chosen from 1 to 120, the
approximate length of the study in days. The current was
then measured on the dates coinciding with the number cho-
sen beginning July 2, 1960.

It must be pointed out that the currents as measured
in this study are not instantaneous, but are taken over a
period of time (Welch, l9h8). Current rate is affected by
a number of well-known variables such as water level, depth
and bottom type,etc. (Longwell, et al., 1932). Since each
shingle is in a different position in the current, differ-
ences in current rate, particularly in the riffle, must
have occurred. The measurement of these differences would
have been extremely difficult and not necessarily useful in
this type of study. McConnell and Sigler (1959) used the
rate of dissolution of standardized salt tablets as an in-
dicator of current rate, but attained correlations only up
to 0.9 meters per second. Since velocities recorded in
this study are much higher than this value, little use
could be found for this type of measurement. Currents may
fluctuate from moment to moment within a given maximum-min-

imum range and even this range can shift a great deal over

12

a short period of a few days as a response to changes in
water level (Blum, 1956). These changes in rate could then
have an effect on the productivity of the aufwuchs on the
shingles. The random sampling of current velocity over a
period of time was an attempt, then, not only to account
for changing water levels and the consequent change in vel-
ocity, but also establishes a tentative maximum-minimum
range for each site.

water temperatures were recorded daily on a thermo-
graph placed on a bridge abutment between the two stations.

The single qualitative study was carried out in Octo-
ber to determine the species composition of the aufwuchs
communities of each situation. The method involved using
the relative frequency of occurrence, 1. e. the number of
times an organism was seen in 50 fields. Before extraction
in alcohol, two shingles, one each from the pool and riffle,
were examined by scraping approximately two square centi-
meters from each shingle onto separate microscOpe slides.

For quantitative determinations of the phytopigment
extract the shingles were brought from the sampling stations
to the laboratory in plastic freezer bags. The shingles
were placed in the freezer compartment of a refrigerator
for twenty-four hours. The aufwuchs were scraped off the
shingle into 95 percent ethanol, using a glass slide-and a
rubber policeman. Freezing facilitated the removal of the
aufwuchs. The bags were then flushed out with ethanol to
remove any aufwuchs which might have been dislodged from

13

Figure 1. Correction graph for phytopigment extracts.

1000

11+

 

I I

I

SOOQF—-

NITS

f

DlOOO—————

KLETT

500.—

 

lOO

 

10

 

Relative Concentration

 

15
the substrate. The use of ethanol rather than acetone pre-
vented the dissolution of the plexiglass, while at the same
time dissolving the phytopigments. The solutions were al-
lowed to stand another twenty-four hours in the dark and
then filtered through glass wool. After filtration the sam-
ples were adjusted to fifty milliliters.

The resulting phytOpigment extract solutions were then
"read” on a Klett-Summerson calorimeter, using a 6h0-700 mu
red filter. Grzenda and Brehmer (1960) found that in only
small concentrations did these phytOpigment solutions fol-
low the theoretical Lambert-Beer Law. As the concentration
of the phytopigments in the 95 percent ethanol increases
the observed and theoretical absorbancy values at 6A0-700
mu become widely divergent. They prepared a graph, using
various concentrations obtained through dilution. The cor-
rection values are obtained from Figure l by reading the
observed Klett units on the ordinate and extending it hori-
zontally to the experimental line. Vertical extension from
the point of interception of the experimental line to the
theoretical Lambert-Beer line and reading of the Klett units
directly perpendicular to it gives the adjusted Klett units.
This value is then multiplied by 2 x 103 to convert to ad-
justed absorbancy AA x 103 and to avoid the use of the deci-
mal. The method is more fully described by Grzenda and
Brehmer (1960).

It is obvious from Figure 1 that observed values above

550 Kletts become impossible to interpret and hence all

16

values above this figure in the raw data and accompanying
figures are only of the proper order and are not exact.
These values, when converted to AA x 103 were interpreted as
15,000 . It seems that if the sample solutions were diluted
to the point where the theoretical and experimental lines
are coincident, then multiplication by the dilution factor
should produce the theoretical Lambert-Beer values. The few
times that this was tried resulted in different values than
when using Figure 1, so it was felt that a slight error in
magnitude would be justified in the case of the higher ob-
served values if some indication of order could be shown.
The writer feels, however, that more work should be done on
this dilution method because, at least, in theory, it should
work.

Only seven complete cycles are shown in Appendix III,
but several others were interrupted due to high water in
June, which made the pickup of the substrates impossible and
again in October when the heavy leaf fall tended to accumup
late on the surface of the shingles. Data from such collec-

tions were discarded.

RESULTS AND CONCLUSIONS

Since the current rates in the pool were generally the
same, always less than 1.0 feet per second, the absorbancy
units (AA x 103) for the three sites have been averaged for
each three day period and depicted as one line in Figures
.2-8. The three sites at Station B have distinct differences
in velocity and are depicted as separate lines for each site
in these figures.

With the exception of Figure 6, which depicts Cycle V,
the pool appears to reach a maximum.within the fifteen-day
cycle and in some cases shows a moderate decline between
the twelfth and fifteenth days. In general, they also reach
much lower maxima than do those in the riffle in spite of a
rapid initial increase during the first three days of a
cycle. These facts are attributable to the almost "immedi-
ate" colonization of the pool substrates and the comparative
ineffectiveness of the slow current velocities to sweep away
the metabolic wastes and to bring in nutrients and gases to
the community, hence inhibiting reproduction after the inp
itial colonization. In addition, much clay and silt is de-
posited along with dead organic matter and these particles
occupy space and perhaps cover the community on the sub?
strate. They then act as a shield and prohibit adequate
light penetration to the community. This last effect would
be more inhibitory as the length of time the particular
shingle was in the water increased, since accumulation of

the inorganic and dead organic particles on the shingle

1?

18

Figures 2-8. AA x 103 per square decimeters
for pool and riffle sites. V9;
pool sites averaged: V , V2, 3,
depicted as separate 1 nes.
Cycles I-VII.

19

Figure 2
CYCLE I July 2—17
10000....—
5000...
I ,’13_._.
1000——
; , V

 

 

DAYS

10000,

5000,

10

 

 

CYCLE

 

20

Figure 3
II July 17—August 2

10000 _____ CYCLE

5000___,

1000 .

500

100

10

 

21

Figure A

III

August 2-17

 

DAYS

 

 

10000.

5000,

1000

T T1|11T1‘ 1

 

E

T F1111]

H
O
O

 

 

AA x 103 per 1.u am?

U‘l
O

10‘

 

 

22

Figure 5
CYCLE IV August 17-September l

 

 

17-

DAYS

 

10000

5000

11|1I11l T“

1000

 

500

1 1 ['111T

T
1

Ahwx 103 per 1.A dm2

 

 

CYCLE V

  
    

23

Figure 6
September 3-13

2h

 

Figure 7
CYCLE VI September 18-October 2
10000,...—
C
5000...
V
10001___
500._- V0
a: o
5
:-
A
a
0
‘1' v
M100
9.
H
3 5°

 

 

l

 

10000

5000

1000

M x 103 per 1.4 dm2

1 1111111] 1

111|1111‘

 

25

Figure 8

CYCLE VII October 2-17

 

 

26
would also increase.

The riffle areas, on the other hand, demonstrate a slow
initial growth (except Site 3), but rise rapidly to reach
higher maxima.. The higher current velocities, V1, at Site
1, Station B, sometimes appeared to be inhibitory, reducing
growth rate and reaching a lower minimum. At the other two.
sites, however, the growth rate was generally greater and
deve10pment of higher maxima was apparent. From this study
it might be concluded that the higher currents are inhibi-
tory in some way, and that the Optimum current lies between
1.00 feet per second and 3.00 feet per second. Both Sites
2 and 3, with velocities of V2 and V3 respectively, show
parallel growth rates generally, with Site 3 growing at a
more rapid rate and reaching higher maxima. Figure 3_con-
cerning Cycle II is apparently an anomaly, with each site
showing a leveling off at about the twelfth day. Field
notes for July 29 and July 30 show thathhe stream was ex-
ceptionally turbid, with much vegetative debris, although
water levels were up only one inch. Maximum current veloc--
ities were measured on the afternoon of July 28, reaching
#.55 feet per second at Site 1, Station B. These conditions
undoubtedly arose from the Opening of the dam in William-
ston. The high turbidity reduced the available light, while
the vegetative debris and molar particles caused attritional
losses.-

The absence of a leveling off in the riffle situation

maxima except occasionally at Site 1 indicates that a longer

27

period of time is needed to reach equilibrium levels. Since
this technique involves the measurement of an accumulated
standing crOp over a period Of time, it is felt that even
at Site 1, Station B, which characteristically was lower,
the inhibitory effect which has been attributed to the high-
er velocities is only apparent. During the fall of 1959,
the steel fence posts were pushed over by duck hunters and
were impossible to retrieve due to high water. In the
spring of 1960 the shingles which had remained attached
through the fall and winter (October 8, l959-May 17, 1960)
were recovered. One shingle from each site in the riffle
was extracted and measured on the Klett-Summerson colorime-
ter. The observed values in uncorrected Klett units were
as follows: Site 1, 806; Site 2, 787; Site 3, 819. These
values were not converted to AA x 103 units, but are cer-
tainly of the same order. The writer feels that such val-
ues would be obtained if a longer time were allowed and that
small differences would occur even at Site 1, which would
eventually reach the same maximum standing crop as at the
other two sites. Clarke (19H6) describes the relation be-
tweem standing crOp and rate of growth of pOpulation. Al-
though the rates of production may differ, he shows that the
equilibrium level may eventually be the same over a longer
period of time. At this point no rate of growth can be de-
termined from the magnitude of the standing crop.

This study does not show an equilibrium level except

for the pool. This level appears to be between 550 AA x 103

28
and 600 AA x 103 in the pool. The absence of an equilibrium

level for the riffle may actually present more useful infor-
mation. Past studies using artificial substrates tended to
rely on accumulated standing crOp as an index to productiv-
ity. Although it is true that accumulated standing crOp was
used as a measurement in this study, the rate of growth
(productivity) can be determined from the changes over a
period of three days. By comparison of these growth curves
then, the riffle appears to be more productive than the pool
with larger attained maxima and a more rapid rate of growth.
The grand average graph, Figure 9, shows the aforementioned
to be true.

From Figures 10-16 it can be shown that the transplants
from the pool to the riffle also were affected by the "phys-
iological richness" of the riffle. The values for the
change between the unmatched shingles are derived from the
algebraic difference between the controls used in the mat-
ched pair. The value for the change between the transplant
and the control was arrived at similarly. Both values rep-
resent the changes over the same period. The values Of the
pool change were averaged since they were generally quite
close, and compared to the change which occurred between
the control member of the matched pair and its "transplant-
ed" counterpart for each site.

In the early stages, three to six days, growth increase
of the transplants was generally behind that of the controls

in the pool. 'As the period of accumulation in the pool was

29

Figure 9. Grand average of accumulation of
phytopigments measured as AA x 103
against time at Vb, V1, V2, V3.

100001

5000

1000

10

Figure

T 1 1 ‘171T1

 

 

 

30

 

 

 

Figures 10-16.

31

Changes in accumulation of pgyto-
pi ents measured as AA x 10 per
1. square decimeters after ”trans-
plant". Pool values from average
change between controls over same
three day period. Riffle values
from change between control and
"transplant".

32

 

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39

increased and larger standing crops transplanted, increases
in productivity resulted. Not only was the rate increased,
but also higher maxima were attained. No changes were noted
in community structure in the tranSplants, but this may have
occurred. The effects of "physiological richness“ may over-
come the ability of the species in the pool to make secure
purchase on the substrate. After the whole substrate has
been colonized in the pool, attritional losses may be re-
duced when moved to the riffle or the increased diffusion
gradient may provide more nutrients and gases for increased
reproduction and cellular growth. Blum (1956) notes that
certain algae which cannot live in rapid current in the
spring and fall seasons, and must live in slow water during
these seasons, are able to attain good growth in the riffles
in the summer.

Although some organisms have inherent current demands
as Welch (1952), Blum (1956), and others show, certain or-
ganisms appear to be able to produce increased growth in
the current, but are incapable initially of colonizing a
bare substrate successfully where excess velocities occur.
Once having successfully established themselves in relative-
ly quiet situations however, they are capable of increased
productivity in the riffle environment. This is demonstrat-
ed by the slow or negative growth shown in the transplants
of three and six days from the pool, and the increasing
growth as the age of the tranSplanted community increases.

Since no transplants were made for longer than three days,

1+0

3
it is possible that these organisms from the pool may have

shown only initial rapid growth and might not have sustain-
ed it. Eventually these transplanted organisms would have
been replaced by other organisms which can compete more
successfully.

Even among the transplants from the pool to the riffle,
the effects of different velocities can be seen. Generally,
the increase in production is less in V1 than in the less
rapid currents of V2 and V3., Much of this difference may
have been derived from attritional losses from the trans-
planted pool community. It is possible that productivity
is as high on the V1 tranSplanted shingles as on the others,
but as soon as growth and multiplication occur, the cells
are swept off the substrate.

The one qualitative study made on October 1%, 1960
shows that the communities differed. The organisms found
in the pool were characterized by two diatoms, Navicula
cryptocephala and Sygedra Elna, with the last named species
occurring in 76 percent of the fields examined. In the rif-
fles, the filamentous diatom Melosira s . occurred in 100~
percent of the fields examined. NO other species were seen
in the riffle samples. The presence of S, ulna and,fi. cr -
tocephala throughout the summer as characteristic species
Of the pool community is unknown, but Peters' (1959) study
showed this community to be a dominant for much of the sum-
mer and early fall. Melosira gp,, however, was present all
through the study. This species grew in dense, brown, gel-

#1

atinous masses on all the riffle shingles and was easily
recognized in the field.

There is no reason to believe that the phytopigments
derived from the different species should exhibit any wide
differences. Some differences, both quantitative and qual-
itative, certainly exist, but evidence of these differences
between species is lacking in the literature. Riley (1938)
discusses the pigments of the various classes of algae, but
points out that differences exist not only between classes,
but also may exist within a single organism due to its phys-
iological state. Gardiner (19h3) seems to feel that the
phytopigment extract method depends for its reliability on
the constancy of the ratios of common pigments in different
classes and in the different seasons. Tucker (19%9) estab-
lishes a high correlation between number of phytOplankters
and phytOpigment density in samples containing over 90 per-
cent diatoms. Little published research is available for
phytopigment quantity and/or quality in taxonomic groups
lower than classes. Strickland (1960) lists phytopigments
present in the various classes of.algae and discusses the
problems involved in determining the amount Of pigments
present in the classes. He states that the pigments found
in the various classes have probably evolved before many
orders and genera. All genera encountered in this study
Are of the BacillariOphyceae and any differences in phyto-o
pigments that might exist are considered to be only transi-

tory and not real, but rather due to the age of the algae

#2
involved, inhibition or exhibition by sunlight, or other
ecological factors affecting their physiological state with
regard to their phytOpigments.

Neel (1951) states that greater consumption of nutri-
ents occurs in the rapids than in the pool.' It would follow
that this is true, particularly since the nutrients are in
greater "physiological" abundance.s That is, more individual
molecules are contacted by an individual organism growing
in the riffles over the same period of time. The rate of
consumption then, would depend on the organisms' ability to
utilize these nutrients, the particular biota present, and
their biomass. In the case of the filamentous diatom, M212?
giga, it seems probable that volumes of water essentially
stationary, but in close contact with.moving water are en-
closed within the filaments. This moving water then can re-
new the supplies of nutrients and gases. Although Ruttner's
"physiological richness" appears to be true, it does not Of-
fer a real eXplanation of the differences exhibited. Blum
(1953) in his work on the Saline River in Michigan, states
that no differences in oxygen content between riffles and
pools were detected either day or night. It has not been
demonstrated that differences in nutrient or gas levels ex-
ist between pool and riffle situations, but it seems obvious
that over a given period of time more nutrients and gases
are presented to a given community of algae in a riffle.
Hence if they have the ability to utilize the nutrients in

reproduction and growth, then the riffle should be more

1+3

productive. Whitford (1960) feels that the difference is a
physical one and that Ruttner's discussion of "physiological
richness" is correct, but does not go far enough. Ferrell,
et a1. (1955) used both organic and inorganic molecules to
show an increased diffusion gradient in water velocities
greater than 0.5 feet per second. This current is able to
reduce the distance between the cell wall and the nutrients
involved to less than 0.25 millimeters. The greater this
proximity, the more able the cell is to capture the molecule
for use in its metabolism. It follows from Whitford's
(1960) study, then, that the higher the current velocity,
the higher the diffusion gradient, and hence increased
growth. This study does not demonstrate this and the writer
feels that excessive velocity might cause greater attrition-
al losses in spite of increased growth rate, but that at
some time their equilibrium level might be similar..

Another ecological factor which might have produced the
better growth of the riffle aufwuchs are differences in
light quality. No evidence can be presented concerning the
exact amount of light penetrating to the substrates in the
pool or riffle. Both received approximately the same amount
and time of shading. Butcher (l9h6) noted that the number
of algae appearing on his submersed slides was always great-
er when the amount of sunlight was greater. Both stations
received the full noon sun for two to three hours through-
out the summer. More exact measurements might have shown

differences in light intensity, but the difference in growth

Ah

rate cannot be attributed only to light quality.

Other ecological factors such as turbidity, depth and
water temperature offer little explanation for the differ-
ences exhibited. The depth at both stations was essentially
the same and all shingles were placed at the same depth from
the surface (0.8). The proximity of the stations rules out
major differences in either water temperature or turbidity.
Peters (1959) found temperatures at a station upstream to
vary less than one degree from that on the thermograph po-
sitioned between the two stations of this study.

The writer feels that if the length of the cycles were
increased and "tranSplants" made from the riffle to the
pool, a more complete and useful study might result. It
seems clear, however, that any future studies using artifi-
cial substrates as a device for gaining information on pro-
ductivity should take into account current velocities which

exceed 1.0 feet per second.

l.

2.

3.

5.

SUMMARY
Differences in growth rate and attained maxima between
aufwuchs communities in pool and riffle situations on
artificial substrates were demonstrated. The riffle, in
general, had a faster growth rate and attained higher
maxima than the pool.
Aufwuchs communities grown in the pool for 3, 6, 9, or
12 days were transplanted to the riffle for three days.
Only slight or negative growth were observed from the
3 and 6 day pool-grown transplants. The 9 and 12 day
transplants showed greatly increased growth after three
days in the current.
Differences in growth rate cannot be attributed to dif-
ferences in gas content, nutrients, turbidity, depth,
temperature or community structure. Light quality and
intensity may have varied, but both received approxi-
mately the same amount of shading and full sun.
Differences exhibited are, therefore, attributed to the
effects of a current with its attendant "physiological
richness" and to an increased diffusion gradient between
the diatom cell walls and a particular nutrient or gas
molecule, which puts this molecule closer to the cell
and as a consequence, becomes more available for meta-
bolic use.
Any future studies using artificial substrates for pro-
ductivity studies should take into account the presence

or absence of a current above 1.0 feet per second.

#5

’+6

Appendix I

Matched pairs tests
Test 1 - Six days
Test 2 - Nine days

1+7

d d2 2
(R-L) (R-L)

8 6%
- h 16
o o
h 16
n 16
- 3 9
19 361
- 1 1
6 36
- 2 u
-1u 256
-10 100
8 6h
15 225
k 16
_ 168E

RIGHT LEFT
85 77
86 90
103 103
10% 100
86 82
122 125
110 91
105 106
113 107
11% 116
12% 138
89 99
inc 132
1M1 126
110 106
_ _ g; =2.267
‘n
612 _ 108k _76. 267
fif""l3””‘

t _‘cT.0_2.267-0 ._ .

 

No significant difference at five per cent

level.

SIX BAX TEST

AAxlO3
RIGHT LEFT
230 226
220 222
229 221
258 2H9
283 253
287 228
229 23%
222 229
226 218
216 222

.....,_' r-

,J. ..
2. 21

#8

.“.~8oo—o_: 0.616

 

d d2
(R-L) (R-L)2
u 16
- 2 u

8 6H

9 81
-10 100
19 361
- s 25
- 7 1+9

8 an
‘88 83%

s- -<82.(:d)2 800- 32.8

 

No significant difference at 5 per cent level.

NINE DAY TEST

49

APPENDIX II--CURRENT DATA

Average

Station Velocity Velocity

and Site Side ft./sec. For Site ft./sec. Date
EB g Ejég n.17 E388 3’
EB E 3:8 . 3.06
21 E 8:;68 O 519
3A E 82323 O 573
2. a 2:528 “65
n L h. 5 8.41 July 13,
2B R 2.68 1960
n L 2.63 2.66
21 E 8322 M32
3A E 82%;? O 572
IE E E13§1 0.590 July 16,
.. L 1,. 5 11.110 1960
EA E 813E7 1.31
31 E giggE‘ 0'711
" L 0.638 0.629
EA E 8:323 0.816
EB if E35 11.87 E3836 28’
33 2 3:12 3.17
3B E EIZE 1-70
EA E 8;;fié 0.750
EA E 8:2EE o 828
EA E 8:322 o 856

50

APPENDIX II--CURRENT DATA, Cont.
Average

Station. , Velocity Velocity

and Site Side ft./sec. For Site ft./sec. Date
EB: E 3.3; 3.83 igggst 6,
83 E 3:38 2.87

33 3, 833 3-36

33 3;: 8:33? 0531

33 3 88% 0-585

.3 3 8:232 0.627

" L 1379 1‘83

93 E 828% 1.37

8A E 8:888- 0~313

E: E §:;1§ 0.523

IB 8 3.5 5 0.5%8 August 2%,
n L 3.06 3'05 1960

33 3 393

8B E 1:33 1.30

88 E glféfi 0.191

58 E 0.361 0'391

3A E 8: 88E 0~382

3. 3 3323' 2'73

n L 1. 71 1.70

33 I; 33333 0.3..

EA E 8 888 0.60%

8A E 8:2E: 0.631

51

APPENDIX II--CURRENT DATA, Cont.

' Average
Station Velocity Velocity
and Site Side ft./sec.

P 8 888 3.10
3 3:33
83 E }:33 1.91
33 3 8:233 0.625
33 3 3:23:- 0....
i: g §E§§8 0.683
" L 3.0M 3'17
53 8 8:83 1:91
83 E 81:; 1.30
%A E 8:3é2 0.335
5A i 8.339 0.371
gA E 0.30& 0.396

0.388

For Site ft./sec. Date

September 22,
1960

October 1,
1960

52

Appendix.III

PhytOpigment extract data

 

53

Table I. Changes in phytopi ent extract
measured as AA x 10 between con-
trol member of matched pairs from
pool. Includes change in AA x 103,
percent change, and logarithm of
percent change.

5a

Table I
CYCLE I
Change 3
Days Site 0 % Change Log % Change
1 81 8100 3.008u9
0-3 2 103 10300 .0128h
3 97 9700 3.98677
1 8 120.0 2. 07918
3-6 2 2 78.8 1. .89653
3 56 57.1 1 7566
1 88 h,.0 1.69020
6.9 2 109 5 .6 1.76790
3 101 6 .5 1. 81623
1 252 9 .0 1. 97 13
9-12 2 305 103.3 2. 01 10
3 315' 162.7 2. .211E:
1 9 1.7 0. 2305
12-15 2 3 8.8 W9
3 " 7 " 1+ 0-9 0.906
CYCLE II
1 111 11100 a. 08532
0.3 2 117 11700 M. 06819
3 107 Iogoo M. 02938
1 110 82 1. 99211
3-6 2 100 n. 7 1.92788
3 92 85.11.92993
1 fig .16.6 1.22011
6-9 2 1 86.2 1.93551
3 137 68. 1. 83569
1 267 6%. 1.80889
9-12 2 23% 57.6 1. 76042
3 192 56.9 1.75511
1 - 12 - 1.9 0.27875
12—15 2 -152 - 23.7 1.37375
3 2n .7 0.67210
CYCLE III
1 97 9700 3% 38677
0-3 2 73 7300 mg
3 76 7600 3. 880 1
1 110 112.2 2. 08999
3-6 2 128 167.5 2.22801
3 112 137.3 2.16820
1 #7 22.5 1. 3 218
5-9 2 17% 87.8 1. 9 389
3 230 122.3 2 08783

55

Table I Cont.
CYCLE III Cont.

Change
Days Site AAx103 % Change Log % Change

1 330 129.9 2.111
9-12 2 2 68.2 1.83338
3 3 9 83.h 1.92117
1 -101 - 17.2 1.23553
12-15 2 - #9 - 7.8 0.89209
3 -106 - 11.9 1.17319

CYCLE IV

1 81 8100 3.90899
0-3 2 67 6700 3.82607
3 93 9 00 3.968u8
1 40 u .7 1.68753
3-6 2 #2 61.7 1.79029
1 1g6 152.8 E'ZZ”52

- . . 2
6-9 2 88 0.0 1.90 83
3 32 21.3 1.32 8
l 227 7 .5 1. 9h 7
9-12 2 217 109.5 2.03991
3 1M9 112.3 2.07628

- - . 0 790
12-15 2 38 11.5 1.06070
3 75 22.6 1.35%11

CYCLE v

1 M5 #500 3.6 321
0‘3 2 55 5500 3.7 036
» g 1 5100 3.70557
6 208.6 2.31 31
3-6 2 122 217.8 2.33 06
3 112 215.3 2.3330h
7h 52.1 1.7168h
6-9 2 92 117. 2.07151
3 26 15. 1.19866
2 316 196.2 2.16H95
9-12 13 78.2 1.89321
1 236 §3°1 l°g163t

. 1. 99
12-15 2 112 79.8 1.90200
3 167 57.» 1.75891

9-12

12-15

9-12

12-15

Site

LUNHle-‘WNHUJNHUJNH

UJMHWNHWNHWNHWNH

56

Table I Cont.

CYCLE VI
Change
AAx103 7 Change

39 3900
91 9100

9 9300
53 13 .0
96 10 .3
82 87.2
237 126.0
252 13H.0
176 100.0
61 18.h
13 11.8
12.5

6 1.5

13 2.8
15 3.7

CYCLE VII

77 700
71 100
$3 6500
56.h

20 27.7
125 102.”
116 126.0
165 175.5
297 120.2
257 123-5
326 125.8
- “'6 801*
" 21 I+05
2 0.39

Log % Change

3.59106
3.9590h
3.968%8
2.1303

2.0182

1.99052
2.10037
2.12710
2.00000
1.26882
1.07188
1.09691
0.17609
O.HH716
0.56820

3.88619
3.85126
3.81291
1.7 28
1. 2M8
1.62737
2.01030
2.10037
2.29928
2.07990
2.09167
2.09968
0.92928
0.65321

.1 oh7712

Table II o-

57

Change in phytopigment extract
between riffle ontrols meas-
ured as AA x 10 . Controls av-
eraged for each site and dgy.

Includes change in AA x 10
percent change, and logarithm

of percent change.

3-6

6-9

9-12

12-15

0-3

6-9

9-12

12-15

Site

WNHle-JWNHUJNHWNH

WNHWNHWNHWNHWNH

WNHWNHWNH

58

Table II
CYCLE I
Change3
AAxlO. % Change
6 600
9 900
#0 #000
2% 3h2.8
2 320.0
7 11h.2
31 100
13% 319.0
179 203.h
136 219.3
172 97.7
3067 1199
310 156.5
1767 507.7
165 . n.90
CYCLE II
n 1100
18 1800
118 11800
#6 92.0
#8 252.6
269 226.0
73 183.1
213 317.9
2 0 59.2
2 H 222.0
782 233.0
1262 2 .2
7 9.0
6 11.8
995 50.2
CYCLE III
1 50
16 1600
13 1300
10 goo
15 8 .2
86 619.2
2% 200
66 206.2
#59 859.0

Log % Change

2.05767
2.00000
2.50 79
2.30 3

2.3910

1.98989
3.059791
2.19851
2.70561
0.69020

2.60206
3.25527

.03188
1.9 379
2.90293
2.35811
2.15568
2.33229
1. 2 2
2. 598%
2. 1996
2.31006

0.9 2%
0.63315
1.70070

3.2OH12

3.%lg9$
2. 9

1.92587
2.78831
2.30103

2.31h29
2.66181

Days

9-12

12-15

9-12

12—15

9-12

12-15

Site

UJNl—‘UJIUI-J

WNHWNHWMHWNHWNH

00:01-4memememe

Table II

59
Cont.

CYCLE III Cont.

Chang 363
AAxlO % Change
20 580.
31 320.
3269 589. 7
210 85. 7
9073 988.5
9995 109.5
CYCLE Iv
11 1100
12 1200
119 11900
25 208.
179 137
1189 1030
106 286. 9
1985 7;a.9
711 7
29g H 1727
1. 0
1995 79.3
501 128.
6995 909.7
7920 225.9
CYCLE v
o 0.00
18 1800
53 5300
51 5100
176 926. 3
795 1380
£5 67 3
11 0 589. 6
2680 335. 9
122 158. 9
2899 213. 9
2996 71.7
269 126.3
10816 258. 5
9025 151. 0

Log % Change

2.76380
2.50569
2. .76693
1.9329

2. 99998
2.03991

3.09133
.0791
.05690
E 31369
13
9.01283
2. 95697
2. 88890
1. 73799
2.23729
0.00000
1.87099
2.10857
2.612 7
2.35392

9-12

12-15

Site

WNHWNmel—JWNHWNH

WNHWNI—‘WNHWNHLUNH

60

Table II Cont .-

CYCLE VI
Change3
AAxlO % Change
10 1000
18 1800
63 6300
25 227.3
19 100.0
39 196.8
0 111.1
195 513.1
2990 1576
65 85.5
666 1573
302 313.5
55 9.0
1666 2.
1300 ‘ 11.
CYCLE v11
6 600
12 1200
17 1700
12 171.9
38 292.3
50 277.7
129 652.6
305 598.0
39 586.7
17 121.6
619 172.9
1063 227.6
551 17g.8
7105 73 .9
8180 539.6

Log % Change

3.00000
3-25527
3.79939
2.35660
2.00000
2.16673
2.09571
2.71020
3-19756
318%?"
2.9962?»
1.59106
1.6309

1.0718

2.77815
3.07918
.23095
2.23901
2.93533
2.

. E65
2.77670
2.76892
2.0899
2.236
2.3 717
2.2 05
2.86975
2.72803

LU

61

Table III. Change in ghytopigment measured
as AA x 10 between control from
pool and transplant from pool to
r1 fle. Includes change in AA x.
10 , percent change,—logarithm
of percent change.

Days

3-6

6-9

9-12

12-15

9-12

12-15

3-6

9-12

12-15

Site

WNHWNHWNHWNH

WNHWNHWNHWNH

WNHWNHWNHWNH

62

Table III
CYCLE I
Change
AAxlO3 % Change
70 85.3
18 17.3
$3 381’;
289 152.6
338 219.
135 50.
53 311-2
7 .
183 28.8
1 1.
79 130.9
CYCLE II
8 7.1
19 11.8
30 27.
a3 19.8
1 g 89.8
21 109.0
186 71.8
189 96.5
591 175.3
59 11.2
72 11.2
396 65.9
CYCLE III
5% 53.0
62.1
66 86.8
97 22.5
232 117.1
230 122.3
195 76.
308 82.7
612 196.9
12 21.7
79 127.9

2311 301.3

Log % Change

1.93095
1.23805
1.00860
1.58933
2.1835

- 2.391

1.702 3
2.060 2
2. 3 2
1. 5939
2.11826
2.11528

0.85126
1.07188
1.99298
1.17026
1.92890
2.03793
1.66795
2.29378
1.09961
1.09961
1.81558

1.72928
1.79 09
1.93 52
1.35218
2.068 6
2.087 3
1.88309
1.917

2.16559
1.33696
1.10517
2.97900

Days

6-9

9-12

12-15

3-6

6-9

9-12

12-15

3-6

9-12

12—15

Site

WNerUHUJNP-JUJNH WNHWNHWNHU’NH

WNHWNHWNHWNH

63

Table III Cont.

CYCLE IV
Chang 883
AAxlO % Change
120 196.0
58 §E.0
333 3 .0 .
277 227.1
350 318.1
107 37 0
297 129.7
898 993$
39
715 172.2
869 262. 5
CYCLE V
32 62.5
201 35 .9
207 398.0
3% 85:2:
99 26. 8
19 6.9
- 100 - 3700
202 106.3
128 29.0
369 9O 3
588 125.7
CYCLE VI
26 65.0
- 60 - 65.2
"' 20 " 21.2
70 79.9
228 122.7
- 16 .- 9:8
10 2.2
890 252.8
- 6 - 16.3
- 138 " 3003
832 210.1

Log % Change

2.169 5
1. 9292
2. 59900
2. 35603
2. 50293
1.56820
2.09 8%
2.69 0
0.81291
2. 23603
2. 91913

1.95569
2.09939

1.81291
1. 8192
1 263
1. 7157
1.10380
2.11227
0.68129
0.9292
.0106
1. 21219
1.98199
2.32293

Lb.- .

-J-PA

Q”) 1

Ir :wu-ung- A-

69
Table III Cont.

CYCLE VII
Change3

Days Site AAxlO % Change Log % Change
1 "' 60 - 7609 - 1088 93
3-6 2 - ,‘I'8 - 66.7 1.82 13
3 39 51.5 1.71181
6-9 2 96 50.0 1.69897
3 99 100.0 2.00000
9-12 2 160 76.9 1.88593
1 - 18 - 3.3 0.51851
12-15 2 95 127.9 2.09687
3 03 137.2 2.13735

‘ Table IV 0

Raw data.

65

Conversion to AA x 103

from Klett units as determ ned
from Figure 1 times 2 x 10 .

‘2
E
E
E
E
y':

CYCLE I July 2-17, 1960'

Days
0-3

6-9

9-12

Code

No.

11a
21a
31a
11b
12b
21b
22b
31b
32b

13a
23a
33a
12a*
22a*
32a*

EEE
33%
33%

15a
25a
3 a
1 a*
29a*
39a*
15b
16b
25b
26b
35b
36b

173
27a
37a
16a*
26a*
36a*
1 b
1 b
27b
28b
3 b
3 b

Kletts

91
33

2

AAx103

82
109
98
6

8
12

3550

66

Table IV

CYCLE I Cont.

Days

9-12

Code

No.

19a
20a
29a
38a
3 a
90a
18a*
28a*
38a*
19b
20b
29b
30b
9b
0b

Kletts

205
197
211
218
219
227
226
309
316
199
200

AAx103

0
E38

685

620

600

665

670
1388
1

96
28
2 ,
1880
3328
3670

CYCLE 11 July 17-Aug. 2, 1960

0-3

3-6

6-9

12a
22a
32a
,11b
12b
21b
22b
31b
32b

19a
29a
39a

11a*
21a*
31a*

EB?
33%
33%

12b,
22b,
32b.

16a
26a
36a

56

33

2

8

11
58
61

111
109
100
60
66
69
22
29
38
31
169
170
o

1

5

121
172
159

112
118
108
a

6
16
22

116
122

259
906

337

 

67
Table IV Cont.

CYCLE II Cont. CYCLE III Aug 2-17, 1960

Code

Code

 

 

Days No. Kletts AAXlO3 Days No. Kletts AAx103
6-9 l3a* ]_ 2 0-3 11a 99 98
238* 12; 965 21a 37 79
338* 17 918 312 38 76
15b 66 132 11b 1 2
16b 38 116 12b ]_ 2
25b 1 l 299 21b 1 2
26b 130 265 22b 16 32
35b 212 590 31b 2 1+
36b 223 695 32b 12 29
lfib' 16+ 18 8
2 bu 1 2 -6 l 2 10‘4- 20
39b, 22 99 3 333a 9 198
33a 9 188
9-12 18a 201 526 122* 75 150
28a 222 690 223* 60 120
383 202 29 32a* 71 192 .3
15a* 183 5 lab -- --- j
2a; :2 333 1b 6 12
'3 a* 2 b 23
17b 182 990 23b 9 18
18b 169 376 33b 30 60
27b 278 1080 3 b 70 190
28b 266 968 11b, 0 0
37b 332 1780 21b. 9 8
38b 3 2 1980 31b. 9 8
16b. 1 2
26bI 9 18 6-9 158 125 255
36b, 16 32 252 163 972
35a 176 18
12-15 19a 200 520 19a* 125 255
20a 197 508 29a* 180 930
29a 199 16 39a* 176 918
302 186 60 15b 10 20
9a 209 599 16b 26 52
0a 207 562 25b 93 86
17a* 211 585 26b 55 110
27a* 239 312 35b 197 19
372* 255 73 36b 297 09
19b 191 98 13b. a 6
20b 132 906 23b. 8
29b 2 2 1120 33b' 5 10
30b 2g2 1020
9b 3 2 2850 9-12 17a 21 585
0b 380 2800 278 21 626
18b. 2 9 378 291 767
28b| 11 22 16a* 189 950
38b. 32 69» 26a* 230 680

Days
9-12

12-15

68

Table IV Cont.

CYCLE III Cont.

Code
No. Kletts AAXIO3
36a* 273 1030
13b 112 226
1 b 129 253
27b 109 218
28b 215 605
37b 922 3880
38b 925 3775
15b. 1 2
25bl 7 19
35b: 36 72
19a 203 38
20a 180 330
292 216 610
302 209 599
39a 211 585
906 237 736
182* 239 712
28a* 30? 1929
38a* 392 3078
19b 195 500
20b 297 809
29b 97 3170
30b 67 5800
9b 510 8015
0b 518 8050
17b, 2 9
27b: 3 6
37b. 1 28

CYCLE IV Aug 17-Sept 1, 1960

0-3

12a 91
22a 9
328 7
11b 6
12b 6
21b 6
22b 7
31b. '35
32b 80
19a 61
29a 55
39a 75
1181"l 101

21a* 63

Days

3-6

9-12

12-15

Code
No.

81*
l b
2:
3E}:
12b

22b
32b

16a
26a
36a
13a*
23a*

26b,
36b:

19a
20a
29a
302
39a

CYCLE IV Cont.

Kletts

1;;
15
73

116

258

326

1

10
21

AAX103.

”El
132
237

902
1696
2

2O
92

 

.1

CYCLE IV Cont.

Code

69

Table IV Cont.

Code

CYCLE V Cont.

 

Days No. Kletts AAxlO3 Days N0. Kletts AAxlO3
12-15 90a 171 903 6-9 39a* 109 208
17a* 205 550 15b 60 120
27a* 283 1130 16b 27 59
372* 290 1200 25b 230 680
19b 250 830 26b ‘3‘3 1990
200 269 952 35b 392 3118
29b 510 8580 36b 26 3890
300 520 8700 130. a 5
39b 579 11900 230. 8
90b 556 10950 330. 12 29
18b| 11 22
28b. 19 28 9-12 170 202 32
38b. 92 89 27a 171 0g
37a 167 38
16a* 115 230
CYCLE v Sept 3-18, 1960 262* 85 170
36a* 168 392
0-3 Ila 23 96 17b 107 219
21a 2 56 18b 102 209
31a 26 52 27b 92 3118
110 1 2 280 57 5250
12h 0 0 37b 530 960.0
21b 8 16 38b 360 2350
22b 6 12 15b. 2 9
310 18 36 25b. 8 16
32b 36 72 35b. 8 16
3-6 13a 71 192 12-15 192 252 898
23a 89 178 20a 291 767
33a 82 169 292 212 590
123* 39 78 308 log 00
222* 126 257 39a 18 1+£0
32a* 127 233 906 182 0
1 b 27 18a* 226 660
1 b 26 52 28a* 291 767
2 b 75 152 382* 267 976
2 b 116 237 19b 180 930
3 b 219 600 200 199 516
3 b 268 989 29b 600 15000
11b: 9 8 300 629 15000
21b. 9 8 9b 638 15000
31b: 29 98 0b 636 15000
6-9 17b, 9 8
15a 108 216 27b. 6 12
25a 133 270 37b. 21 92
33a 95 190
l a* 115 2&5
29a* 122 2 9

70

Table IV Cont. '

CYCLE VI Sept 18-Oct 2, 1960

Days
0-3

3-6

6-9

9-12

Code
No.

112
212
312
11b
12b
21b
22b
31b
32b

132
23a

33a
122*

33b.
17a
16a*

262*
362*

Kletts

293

AAX103

352
169
212

62
90

368
1696
3600

16
l6
16

92

396
15
50

12 2

Days
9-12

12-15

CYCLE VII Oct 2-Oct 17, 1960

0-3

Code
No.

l b
18b
27b
28b
37b
38b
15b.
25b|
35b:

19a
20a
29a
308
9a
0a
182*
282*
38a*
19b
20b
29b
30b
9b

128
22a
32a
11b
12b
21b
22b
31b
32b

19a

29a ,

39a

lla*
2la*
312*

CYCLE VI Cont.

Kletts
79
62

283
2
556
556

8
12

170
169
188
187
170
171
2..
292
96
100

92

61
27
73
6
10
9

 

 

.. . .‘u,

71
Table IV Cont.

CYCLE VII Cont. CYCLE VII Cont.

Code 3 Code
Days No. Kletts AAxlO Days No. Kletts
3-6 lab 7 19 12-15 172* 201
1 b 12 29 272* 276
23b 30 60 372* 309
2 b 21 92 19b 266
33b 33 66 20b 291
3 b 35 70 29b 519
12b 7 19 30b 998
22b 7 19 9b 555
32b 15 30 0b 512
18b. 7
6-9 16a 121 29 28b. 11
26a 109 20 38b. 19
36a 127 259
132* 53 106
232* 69 1&8
332* 99 1 8
15b 51 102
160 72 199
25b 187 965
26b 121 297
35b 199 16
36b 176 18
19b. 9 8
29b. 6 .12
34b. 13 26
9-12 18a 209
28a 187 38;
382 211 585
15a* 101 202
25a* 162 368
352* 291 1219
1 b 199 311
1 b 199 323
27b 239 712
28b 292 1228
37b 278 1080
38b 392 1980
16b, 0 o,
26bl 9 8
12-15 19a 198 12
202 191 89
29a 179 927
302 186 960
9a 219 600
0a 209 579

72

LITERATURE CITED

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Clarke, G. L. 1996. Dynamics of production in a marine
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Cooke, W. B. 1956. Colonization of artificial bare areas
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Ferrell, J. K., K. 0.-Beatty, Jr. and F. M. Richardson.
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1 .

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Meehan, W. R. 1958. The distribution and growth of fish
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73

Neel, J. K. 1951. Interrelationships of certain physical
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Odum, E. P. 1953. Fundamentals of ecology. W. B. Saun-
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Odum, H. T. 1956. Primary production in flowing waters.
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Peters, J. C. 1959. An evaluation of the use of artifi-
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duction. Master's Thesis. Michigan State Univer-
SitX-o-

Riley, G. A. 1938. The measurement of phytoplankton.
Int. Rev. d. ges. Hydrobiol. u. Hydrgr. 36: 371-373.

Ruttner, F. 1953. Fundamentals of limnology. Transl.
from German by D. G. Frey and F. E. J. Fry. 292 pp.

Stokes, R. M. 1960. The effects of limiting concentra-
tions of nitrogen on primary production in an artifi-
cial stream. Master's Thesis. Michigan State Uni-
versity.

Strickland, J. D. H. 1960. Measuring the production of
marine phytoplankton. Fish. Res. Bd. Canada. Bull-
etin 122. 172 pp.

Tucker, A. 1999. Phytopigment extraction as a method of
quantitative analysis of phytoplankton. Tr. Am.
Micro. Soc. LXVIII, No. 1. 21-33.

Welch, P. S. 1998. Limnological techni ues. McGraWhHill
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302-309.

 

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