Water Research Pergamon Press 1968. Vol. 2, pp. 311-324.

Primed in Great Britain

CONDENSED PHOSPHATE DEGRADATION AND ASSIMILA-

TION BY SELECTED BLUE-GREEN ALGAE
ERNST M. DAVIS* a n d MAXWELL J. WILCOMBt
* Environmental Health Engineering Laboratories, Civil Engineering Department, The University
of Texas, Austin, Texas U.S.A.
5"Office of Solid Wastes Training, U.S.P.H.S., R. A. Taft Sanitary EngineeringCenter, Cincinnati,
Ohio, U.S.A.

(Received 9 February 1968)

Abstract--Pure cultures of blue-green algae which are found in waste stabilization ponds were
exposed to known quantities of three condensed phosphates. The rates of hydrolysis of the
condensed phosphates were calculated. Information given by the laboratory experimentation
showed that the phosphorus uptake and hydrolysis rates varied among the blue-green genera
and even different species of a single genus. Results were obtained from static culture
chambers and horizontal columns four feet long through which the phosphorus supplemented
medium was pumped over actively growing mats of algae. The test algae used were Anabaena
cylindrica, Anacystis nidulans, Gloeocapsa alpicola, Oseillatoria borneti, O. chalybia, O. formosa,
O. tenuis and Phormidium faveolarum. The test phosphates were sodium tripolyphosphate,
tetrapotassium pyrophosphate, and sodium hexametaphosphate.

OBSERVATIONSon numbers of waste stabilization ponds in the Southwest by PORGES

and MACKENTHAN(1963) and R~ID et al. (1964) have shown that during certain times
of the year members of the Division Cyanophyta (blue-green algae) became the
predominant, if not the only, photosynthetic group in the pond. This establishment
usually began in July and extended well into September. At the same time, analyses
of the waters of the ponds in question showed, on occasion, the effluent total phos-
porus concentrations to be higher than the influent concentration. The questions that
immediately arose were whether the blue-green algae were responsible for this release
and exactly what their role was regarding their capacity for the assimilation of different
forms of phosphates. The condensed phosphates used in this work were those which
are most likely to enter sewage systems and were chosen primarily from the stand-
point of their current use as synthetic detergent builders and water works corrosion
control measures. They are tetrapotassium pyrophosphate (TKPP), sodium tripoly-
phosphate (STPP), and sodium hexametaphosphate (SHMP). This work was an attempt
to determine the condensed phosphate degradation activity and assimilative capacity
of various representative species of blue-green algae for comparison with data already
obtained by DAVIS and WILCOMB(1967) on selected species of green algae. The algal
cultures used were limited strictly to single species and, as far as possible under labora-
tory conditions, were bacteria-free. Cultures of these algae were obtained from the
collection at Indiana University (STARR, 1960), in order to insure the reproducibility
of cultures. They were representative of genera which have been shown by DAVISet al.
(1965) and REID et al. (1964) to occur in waste stabilization lagoons in the Southwest.
The blue-green species used were Anabaena cylindrica, Anacystis nidulans, Gloeocapsa
alpicola, Oscillatoria borneti, O. chalybia, O. formosa, O. tenuis and Phormidium
faveolarum. A discussion of other literature references pertinent to this investigation
has been included in the earlier presentation by DAvis and WILCOMB(1967).
311
312 ERNST M. DAVIS and MAXWELL J. WILCOMB

MATERIALS AND METHODS

Complete descriptions of the nutrient medium in which the algae were cultured,
culture chambers, colorimetric determination of the phosphates, controlled environ-
mental conditions, and other pertinent factors in the experimentation have been
presented earlier by DAVIS and WILCOMB(1967). In order that laboratory error would
be minimal, laboratory conditions were held constant during the phase of the investi-
gation with the blue-green algae as they were when the green algae were tested.

RESULTS
The static experimental runs
FIGURES 1 through 8 present the graphic results of the static runs involving the test
phosphates. They have been corrected for hydrolysis rates in order to gain a more
meaningful insight into the actual mechanisms of phosphate metabolism by the algae
in question. In all cases a point on the corresponding condensed phosphate curve
represents a single membrane filtered sample represented by the analytical components.
Inclusion of four species of Oscillatoria in the work was done to demonstrate the
different activities of specific subgeneric groups.
A comparison of the ortho and condensed curves for Anacystis nidulans showed
assimilation of both during the early stages of growth. It is important to note that an
increase in any condensed phosphate curve, above the concentration indicated at the
start of a run was, in all probability, due to the release of other inorganic condensed
forms or organic forms from the species in question.

12 400
--condensed
ii- ---- -- ortho
-----weight "330
i0-

9- -300

eo
~so ~4
o ?.

6- -200
J
5-
-150 -o
4-

-i00
tD
2--

i- J 50

' '---~ ~ ' ~ ' ' ' I ' ' ' ' I ' ' ' ' ' ' ' ' ' "'I ' ' ' '
5 i0 15 20 25 30
Time (days)

FIG. 1. Effect of Anabaena cylindrica on sodium tripolyphosphate, tetrapotassium pyrophos-

phate, and sodium hexametaphosphate in MSS medium with a typical growth curve for the
organism.
 Condensed Phosphate Degradation and Assimilation by Selected Blue-Green Algae 313
12 400
ii
1 ....
condensed
ortho

-----weight -350
i0
~9-~ _ ~ .3oo

150

3 -4--- - 4- ~ -i00

./I so
' '' ~ ' , I ' ' ' ' I ! , | , I ' ' ' ' I ' ' ' ' I ' ' ' '
5 i0 15 20 25 30
Time (days)

F]o. 2. Effect ofAnacystis nidulanson sodium tripolyphosphate, tetrapotassium pryophosphate,

and sodium hexametaphosphate in MSS medium with a typical growth curve for the organism.

12 condensed
I 400
ii. . . . ortho
----- weight 350
i0-
[ 300

I
~.5.. A
- Ir
~ ~ 150 ~
:~_= _ -_-'~z_~_ . . . . .

3-~ ~ lOO

2-
50

5 i0 15 20 25 30
Time (days)

FIG. 3. Effect of Gloeocapsa alpicola on sodium tripolyphosphate, tetrapotassium pyrophos-

phate, and sodium hexametaphosphate in MSS medium with a typical growth curve for the
organism.

E W
314 ERNST M . DAVIS a n d MAXWELL J . WILCOMB

12
- 400
-" condensed
ii- . . . . ortho
-----weight - 350
i0-

"e-
- 9-
300

~8.
250

~ 6- ~ SHMP _
200 .~t~o

150
4- "~

g i00
""--...
5O
1- / " o

' ' ' ' ' I ' '' ' " I ' ' ' , l ' ' i , I i 'I i i I ' ' ' '
5 i0 15 20 25 30
Time (days)

FIO. 4. Effect of Oscillatoria bornetion sodium tripolyphosphate, tetrapotassiumpyrophosphate,

and sodium hexametaphosphate in MSS medium with a typical growth curve for the organism.

12 "400
condensed
11 ------ortho
-----weight
....7j ' j -350
I0

(9

//"J -300

.25o
eo

,~ 7
O
- - "*- - - 5"
-2oo .~
f ~ ~'~I" I ~"

~ 5
/
"150 ~a
<

~ 3 -I00

-50

...~..m '''/ ^ SHMP

, , l , l 1 1 , , l , ]

5 /o .... ,~ .... ;o .... ;~' 30

Time (days)

Fio. 5. Effect of Oscillatoria chalybia on sodium tripolyphosphate, tetrapotassium pyrophos-

phate, and sodium hexametaphosphate in MSS medium with a typical growth curve for the
organism.
 Condensed Phosphate Degradation and Assimilation by Selected Blue-Green Algae 315
12 400
condensed
11- . . . . ortho
-----weight -350
iCe

~9- -300

SHMP 250

-0

150

5 I0 15 20 25 30
Time (days)

FIG. 6. Effect of Oscillatoria formosa on sodium tripolyphosphate, tetrapotassium pyrophos-

phate, and sodium hexametaphosphate in MSS medium with a typical growth curve for the
organism.

12 400
condensed
ii . . . . . ortho
~--- weight - 350
i0-

~-~ 9- - 300

~0
o
o~
ss~ ~ - 250
o 7- A'"

~" 6- 11..
~'4"'~'~'" -200 ..4

5 ~ ' ~ .~. ~ ~-~-f ~ . ------.I -o

~4

1 ~ 50

5 i0 15 20 25 30
Time (days)

FIG. 7. Effect of Oscillatoria tenuis on sodium tripolyphosphate, tetrapotassium pyrophos-

phate, and sodium hexametaphosphate in MSS medium with a typical growth curve for the
organism.
316 ERNST M. DAVIS and MAXWELL J. WILCOMB
12 400
condensed
ii- . . . . ortho
-----weight 350
10-

300

o~
~o
o~ 250

6- ___ ----
-- 0
0 0
200

~. 5-
150

I00

50

l-

I I ! ' ' + ' I ' ' + ' I l l i l

5 i0 15 20 25 30
Time (days)

FIG.8. Effectof Phormidiumfaveolarumon sodiumtripolyphosphate,tetrapotassiumpyrophos-

phate, and sodium hexametaphosphatein MSS medium with a typical g~owthcurve for the
organism.

Among the filamentous species tested, Anabaena eylindrica (FIG. 1) seemed to

exhibit incorporation of ortho phosphate at about the same rate as it returned conden-
sed phosphate forms to the medium. This indicated a possible preference for the ortho
form over condensed forms. After the first week of culture, slow hydrolysis of all three
condensed forms was evidently accomplished by this organism as well as by Phormi-
dium faveolarum (FIG. 8). Comparison of different species of the genus Oscillatoria
(FIGs. 4-7) showed pronounced degradation of the condensed forms. In almost all cases,
a reduction of the quantities of condensed phosphates occurred over the entire period
of testing. The only exception was the filamentous species, Oscillatoria borneti, in the
presence of sodium tripolyphosphate (FIG. 4).
The growth curves for each species (FIos. 1-8) were calculated on a dry weight basis
and taken from control runs. Data for these will be presented and discussed later in
this presentation.
TABLE 1 gives additional interpretive data of the phosphate curves shown in FIcs.
1-8. The per cent total phosphorus used was calculated on the basis of the total
amount of phosphorus present at the beginning of each run, and this in turn was
considered to be the 100 per cent value. Examination of these data reveals wide
differences in percentages of phosphorus assimilated by the various species. This
would appear to be questionable in so far as constancy among, say, blue-green algae is
concerned but it must be pointed out that the maximum quantities were taken from
the medium at different times during the 30 day testing period. Anacystis nidulans
reached its maximum uptake earlier than any of the other algal species tested. The
 Condensed Phosphate Degradation and Assimilation by Selected Blue-Green Algae 317

i m p o r t a n c e of this d a t u m lies in the fact that after the times indicated, the algae were
r e t u r n i n g p h o s p h a t e c o m p o u n d s to the m e d i u m . This was the case of virtually all the
species which showed a m a x i m u m u p t a k e p r i o r to the e n d o f the test run, or 30 days.
C o n t i n u o u s u p t a k e over the entire 30 day period o f all three phosphates tested only
occurred with Anabaena cylindrica, Gloeocapsa alpicola, Oscillatoria formosa, a n d O.
tenuis. T h e rate c o n s t a n t s were almost always different i n those cases a n d were, with
only one exception, higher for s o d i u m h e x a m e t a p h o s p h a t e u p t a k e t h a n tetrapotassium
p y r o p h o s p h a t e uptake. By c o m p a r i s o n , the rate constants for s o d i u m hexametaphos-
phate were higher, or equal to those of s o d i u m tripolyphosphate, b u t never lower. This

TABLE 1. RELATIONSHIPSOF TOTALPHOSPHORUSUPTAKEDURING

STATICRUNS*

Algal Test Total

species phosphate usedt Day~ k
(%)

Ababaena cylindrica STPP 3.25 30 - 0.010

TKPP 14.38 30 -0.005
SHMP 5.57 30 -0.019

Anacystis nidulans STPP 19.00 12 - 0.009

TKPP 22.85 5 -0.052
SHMP 6.53 5 -0.013

Gloeocapsa alpicola STPP 58.90 30 - 0.028

TKPP 14.88 30 -0.005
SHMP 52.10 30 - 0.028

Oscillatoria borneti STPP 2.51 30 - 0.007

TKPP 60.59 21 - 0.044
SHMP 39.40 25 - 0.020

Oscillatoria chalybia STPP 54.55 10 -0.079

TKPP 2.08 30 - 0.001
SHMP 2.96 30 - 0.001

Oscillatoria formosa STPP 34.44 30 - 0.014

TKPP 10.04 30 -0.003
SHMP 33.83 30 -0.014

Oscillatoria tenuis STPP 2.68 30 - 0.001

TKPP 16.24 30 - 0.006
SHMP 14.75 30 -0.005

Phormidium faveolarum STPP 19.72 20 -0.011

TKPP 23.40 21 -0.013
SHMP 5.51 11 -0.005

* Corresponds to data shown in FIos. 1-8.

t Maximum amount of total P present at t = 0 taken from medium.
Day observation of (I") made, from t = 0.
Given in ha N,/N = - kt units day from t = 0 to day indicated
by (~:).
318 ERNST M. DAVIS and MAXWELL J. WILCOMB

might indicate that hydrolysis of sodium hexametaphosphate by enzymatic action is

more easily accomplished than that of sodium tripolyphosphate or tetrapotassium
pyrophosphate. This is partially substantiated by the relative stability of sodium
hexametaphosphate in the synthetic growth medium as shown earlier by DAVIS and
WILCOMB (1967).
The results of the static control runs are shown in TABLE2. Series of 125-ml Eden-
meyer flasks containing 100 ml of the nutrient medium were used for these tests.
Excepting flask size, the conditions which existed during the static runs and those in the
control runs were identical. The data corresponds to analyses made on the filtrate
taken when weights of the algae were made, as shown on the growth curves (FIGS. 1-8).
In the controls, and without exception, when only ortho phosphate was present at the
beginning of a run, the condensed forms increased during the 30 day 'period of
observation (TABLE 2). Concentrations are indicated for each flask on the same day as
weight measurements are reported.

TABLE 2. PHOSPHATE CONCENTRATIONS IN FILTRATE DURING

CONTROL RUNS*

Phosphate
Algal (mg/1)
species D a y t Ortho Condensed

Anabaenacylindrica 0 4.50 0,02

5 4.01 0.09

10 3.65 0.15
15 3.20 0.19

20 2.99 1.12
25 2.40 1.40
30 2.72 1.42

Anacyst~nidulans 0 4.47 0.01

5 4.01 0.03
10 2.13 0.09
15 3,02 0.62
20 3,50 0.90
25 3,77 0.56
30 3,92 0.21

Gloeocapsa alpicola 0 4,54 0.04

2 3.92 0.04
5 3.41 0.09
10 2.62 0.31
15 2.01 0.46
19 1.64 0.71
25 1.42 0.91
30 1.39 1.61

Oscillatoria borneti 0 4.51 0.01

5 4.45 0.03
8 4.02 0.11
 Condensed Phosphate Degradation and Assimilation by Selected Blue-Green Algae 319
Tab~ 2 cont.

Phosphate
Algal (mg/1)
species Dayt Ortho Condensed

12 3.72 0.15
15 3.51 0.27
20 1.62 0.41
24 1.99 1.12
30 2.97 1.27

Oscillatoria chalybia 0 4.97 0.05

4 4.77 0.09
5 4.75 0.11
10 3.13 0.40
12 2.68 0.40
15 2.01 0.31
19 1.86 0.53
20 1.35 0.86
24 1.94 0.99
25 1.72 1.12
30 1.88 1.35

Oscillatoria formosa 0 3.97 0.02

4 3.90 0.02
8 3.65 0.04
12 3.11 0.09
15 3.00 0.17
20 2.08 0.65
24 2.41 0.71
30 2.49 0.30

Oscillatoria tenuis 0 4.55 0.05

5 4.50 0.05
10 4.38 0.04
15 3.82 0.08
20 3.51 0.13
25 3.69 0.15
30 3.78 0.27

Phormidium faveolorum 0 4.10 0.05

2 4.03 0.05
8 3.76 0.08
11 3.21 0.12
15 2.91 0.16
19 3.07 0.29
28 3.41 0.35
30 3.50 0.44

* 125 ml Erlenmeyer flasks containing 100 ml of medium.

? Indicates same points shown on growth curves, FIGS 1-8.

TABLE 3 shows g r o w t h rate c o n s ta n t s f o r the c o n t r o l runs f o r each species. T h e

c o n s t an t s are at least t e n f o l d those o f p h o s p h o r u s u p t a k e f o r all species reported. T h e
c o n s t an t s are n o t as high as th o s e r e p o r t e d by KRATZ an d MYERS (1955) an d MYERS
320 ERNST M. DAvis a n d MAXWELL J. WILCOMB

(1961), indicating that the nutrient medium as used in this investigation was not a
rapid growth medium but one which permitted optimum growth to occur.
The data related to phosphorus removal in TABLE 3 was derived from those given in
TABLE 2. The m a x i m u m amount of phosphorus taken from the medium from zero
time to the indicated day and the corresponding per cent of the total phosphorus
present at the onset of the control run is related. F r o m these data the uptake rate
constants were calculated. It is immediately obvious that phosphorus uptake among
algae differs a m o n g genera and species and can only be interpreted by examination of
the environment at one particular time.

TABLE 3. CONTROL RUNS DATA SHOWING GROWTH CONSTANTS AND PHOSPHATE

REMOVAL RATE CONSTANTS

Maximum Total P
phosphate present at
Species k* Day removed t= 0 Day1" k:~
(mg/l) (%)

dnabaena cylindrica 0.19 25 1.13 25.0 15 -0.023

dnacystis nidulans 0.24 20 2.26 50.5 10 -0.070
Gloeocapsa alpicola 0.07 30 2.25 49.2 25 - 0.027
Oscillatoria borneti 0.23 23 2.49 55.2 20 - 0.040
Oscillatoria chalybia 0.20 30 2.94 59.6 12 -0.073
Oscillatoriaformosa 0.16 30 1.26 31.6 20 -0.019
Oseillatoria tenuis 0.17 30 0.96 20.8 20 -0.012
Phormidium faveolarum 0.16 30 1.08 26.0 15 - 0.020

* In In N/No = kt units/day on dry wt. basis.

t Related to data in TABLE2.
In In N o / N = - kt units/day.

Comparison of the phosphorus uptake rate constants of TABLE 1 and 3 shows a

rather striking fact. With few individual exceptions, the rates o f phosphorus
uptake were greater when only ortho phosphate was present than when condensed
forms were present. This m a y indicate a slight inhibition of the phosphorylation
mechanism in the algae when a predominance of condensed phosphate is present.
Another possibility, of course, would be that saturation of the luxury uptake mechanism
had occurred.

T h e c o l u m n runs
The same species used in the static runs were used for this phase of the investigation
with the exception of O s c i l l a t o r i a f o r m o s a and O. tenuis. Gross bacterial contamination
o f the cultures prevented these from being used. In all instances sterile conditions were
maintained throughout the experimentation. After each 36 hr run involving a column,
the culture was allowed to stabilize itself for a period of at least 48 hours before another
run was started. TABLES 4 and 5 show the data taken during the column runs. The
 Condensed Phosphate Degradation and Assimilation by Selected Blue-Green Algae 321
expected value f o r p h o s p h o r u s c o n c e n t r a t i o n , shown in TABLE 5, was c a l c u l a t e d f r o m
the e q u a t i o n
d x = 0 . 1 ( x s ) d t - 0 . 1 ( x c ) dt.
T h e n o t a t i o n x, = c o n c e n t r a t i o n o f ortho o r c o n d e n s e d p h o s p h a t e being a d d e d in the
s t o c k n u t r i e n t m e d i u m a n d x , - - - c o n c e n t r a t i o n o f ortho o r c o n d e n s e d p h o s p h a t e
c o n t e n t o f the c o l u m n at a n y time = t. I n t e g r a t i n g at t = 0 where the c o n c e n t r a t i o n o f

TABLE4. ANALYSES OF CONDENSED PHOSPHATES ADDED TO COLUMNS AND COLUMN CONTENTS

AT t=O

Phospate stock analysis Column phosphate content

Column Form (rag/l) (rag/l)
with added Ortho Condensed Ortho Condensed

Anabaena cylindrica STPP 0.09 3.50 3.14 0.00

TKPP 2.02 4.18 3.26 1.37
SHMP 0.39 4.74 1.76 1.83

Anacystis nidulans STPP 0.06 6.01 0.01 0.24

TKPP 0.46 4.37 1.14 0.74
SHMP 0.13 5.34 1.66 0.01

Gloeocapsa alpicola STPP 0.00 4.63 0.07 0.13

TKPP 0.07 4.77 0.59 0.02
SHMP 0.65 7.85 0.62 1.25

Oscillatoria borneti STPP 0.01 6.16 0.36 2.13

TKPP 0.05 4.20 0.63 1.46
SHMP 0.48 3.97 0.26 0.17

Oscillatoria chalybia STPP 0.05 6.05 0.01 0.24

TKPP 0.06 5.94 0.88 0.78
SHMP 0.03 5.55 0.50 0.55

Phormidium faveolarum STPP 0.01 5.66 0.01 1.42

TKPP 0.03 4.30 0.01 0.09
SHMP 0.09 4.86 0.17 0.26

Control STPP 0.06 6.14 6.00 0.38

TKPP 0.04 4.83 4.25 0.36
SHMP 0.03 4.28 4.49 0.10
STPP 0.02 5.04 4.20 0.09
TKPP 0.03 4.83 4.52 0.10
SHMP 0.03 5.11 4.55 0.39

stock n u t r i e n t (xs) in the c o l u m n was zero allowed s o l u t i o n o f the c o n s t a n t . T h e n the

expected value at t = 36 h r (1.5 days) was o b t a i n e d . D u p l i c a t e c o n t r o l r u n s (TABLES 4
a n d 5) with the three f o r m s o f c o n d e n s e d p h o s p h a t e s were r u n to o b t a i n average
figures f o r r e d u c t i o n f r o m t h e o r e t i c a l expected values to a c t u a l a s s a y values. C o m p a r i -
son o f the expected values w i t h those o b t a i n e d a t 36 h r shows surprisingly little
p h o s p h o r u s t a k e n f r o m the m e d i u m b y s o r p t i o n a n d / o r a s s i m i l a t i o n m e c h a n i s m s . T h e
322 ERNST M. DAVIS a n d MAXWELL J. WILCOMB

percentages o f phosphorus assimilated during the control runs were: 5.5 Yo, STPP;
4.5 ~o, T K P P ; 5.5 Yo, SHMP. In all citations throughout the text, the concentrations
of phosphorus are as phosphorus and not phosphate (PO4).
TABLE 6 is a compilation of the data from TABLES4 and 5. The rate constants for
phosphorus uptake shown indicate the differences of the 36 hr content analyses and

TABLE 5. ANALYSES OF COLUMN CONTENTS AT 36 HR AND EXPECTED VALUES OF CONTENTS

Phosphate content analysis Expectedphosphate values

Column Form (mg/1) (nag/l)
with added Ortho Condensed Ortho Condensed

Anabaena cylindrica STPP 2.74 0.72 0.17 3.41

TKPP 2.35 1.50 2.05 4.11
SHMP 3.26 1.37 0.40 4.66

Anacystis nidulans STPP 0.72 0.46 0.06 5.85

TKPP 3.07 0.13 0.48 4.27
SHMP 1.96 0.68 0.17 5.20

Gloeocapsa alpicola STPP 1.89 0.14 0,00 4.55

TKPP 2.38 0.16 0,08 4.64
SHMP 3.46 0.38 0,65 7.67

Oscillatoria borneti STPP 0.60 4.66 0.02 6.05

TKPP 0.38 2.34 0.07 4.13
SHMP 1.04 1.08 0.47 3.87

Oscillatoria chalybia STPP 0.46 1.76 0.05 5.89

TKPP 0.14 2.08 0.08 5.80
SHMP 0.47 1.96 0.04 5.41

Phormidium faveolarum STPP 0.07 0.07 0.01 5.55

TKPP 0.80 0.24 0.03 4.19
SHMP 0.89 0.93 0.09 4.74

Control STPP 0.18 5.72 0.22 5.98

TKPP 0.09 4.57 0.15 4.71
SHMP 0.12 3.95 1.15 4.17
STPP 0.11 4.80 0.13 4.91
TKPP 0.05 4.49 0. I 1 4.65
SHMP 0.09 4.61 0.14 4.83

the expected values at 36 hours corrected for sorption rates. Without exception, the
uptake rate constants for all the blue-green algae in the columns were much higher than
those for the static runs. These data indicate that a nutrient medium passing over
actively growing mats of algae has a much greater potential for being reduced in
phosphorus concentration than it does in a static situation. The ortho phosphate content
analysis column in TABLE 5 shows that in all instances (except, of course, the controls)
 Condensed Phosphate Degradation and Assimilation by Selected Blue-Green Algae 323
the ortho phosphate values were higher than the expected values. This can only mean
a higher rate of hydrolysis from the condensed forms to the ortho phosphate form in
comparison to a static situation.

TABLE6. RATE CONSTANTS FOR TOTAL PHOSPHORUS TAKEN FROM MSS

MEDIUM DURING COLUMN RUNS*

Column with Form added kt Total used:~

(%)

Anabaena cylindrica STPP -0.019 2.95

TKPP -0.310 37.48
SHMP - 0.069 8.50

Anaeystis nidulans STPP - 1.075 80.20

TKPP - 0.267 32.60
SHMP - 0.472 50.00

Gloeocapsa alpicola STPP -0.537 55.40

TKPP - 0.413 46.20
SHMP -0.513 53.70

Oscillatoria borneti STPP -0.093 13.25

TKPP -0.288 35.20
SHMP - 0.475 51.20

Oscillatoria chalybia STPP - 0.655 62.60

TKPP -0.650 62.30
SHMP -0.537 55.40

Phormidium faveolarum STPP -0.916 97.49

TKPP - 0.932 75.40
SHMP --0.650 62.30

* Corrected for sorption losses in 36 hr.

t In No~N= --kt units/day.
:~Per cent total phosphorus taken from medium in 36 hr.

DISCUSSION
These studies of blue-green algae phosphate uptake and hydrolysis activities, studied
by means of static cultures and nutrient batch fed cultures in horizontal columns
indicated a high degree of correlation within each type of experiment but little simi-
larity between the two types regarding the magnitude of the rates of uptake and hydroly-
sis of the phosphorus. At least one blue-green algae species tested showed preference
for ortho phosphate over condensed phosphate forms for its metabolism. Condensed
inorganic and/or organic forms of phosphate may be returned to the aqueous environ-
ment during certain growth phases of blue-green species. Anabaena cylindrica,
Gloeocapsa alpicola, Oscillatoria formosa, and O. tenuis were the only species which
showed continuous uptake of sodium tripolyphosphate, tetrapotassium pyrophos-
phate, and sodium hexametaphosphate over the 30 day test periods in static culture.
324 ERNST M. DAWSand MAXWELLJ. WILCOMB
The average rate constants for total phosphate uptake of all species tested in static
conditions were -0.019, - 0 . 0 1 6 and - 0 . 0 1 2 with STPP, TKPP, and SHMP respect-
ively. Even though the rate for SHMP was lower than the other two, enzymatic
hydrolysis of the meta form may be more easily accomplished. All algal species tested
were shown to return some amounts of condensed phosphates to the medium during
the latter stages of their growth cycles. This may account for occasional lagoon
effluents.
The average growth rate of the test algal species was 0.177, on the order of 10 times
the phosphate uptake rates. The average total phosphorus uptake rate constants for
all algal species tested in the columns were -0.549, -0.476, and - 0 . 4 5 2 for STPP,
TKPP, and SHMP respectively. These correspond to 28.9, 29.7, and 37.6 times those
values for static uptake rates. Considering the times involved (30 days for the static
tests vs. 1.5 days in columns) this clearly demonstrates that phosphorus had a much
greater potential for removal when passing over actively growing algal mats than when
in a static environment.

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