Resistance and susceptibility of algae to decomposition

by natural microbial communities

Douglas GunnisonL and Martin Alexander
Graduate Field of Ecology and Evolutionary Biology,
Cornell University, Ithaca, New York 14850

Abstract
The susceptibility to microbial decomposition of species of 14 algae was assessed in
pond water and with inocula from several environments. Some of the algae were destroyed
in short periods, but others withstood microbial digestion for more than 4 weeks. The
production of toxins did not account for the resistance of those algae not readily destroyed
microbiologically. The suitability of the cell walls as a substrate for microorganisms was
correlated with the longevity of three susceptible ( ChZamydomonas reinhardtii, CyZindro-
spermum sp., and Ulothrix fimbrata) and three resistant (Fischerella muscicola, Pediastrum
duplex, and Staurastrum sp.) algae. Microorganisms able to digest walls of the susceptible
but not of the resistant algae wcrc also isolated. The differing susceptibilities to decom-
position may be related to the relative biodegradabilities of specific components of the
algal walls. -

Although algal growth can be controlled
or modified by chemical means and by
reducing nutrient inputs into waters, little
is known of the biological control mecha-
nisms that limit algal growth under normal
conditions or cause the sudden termination
of aquatic blooms. This study was under-
taken to gain an understanding of some of
the biochemical factors that may be impor-
tant in programs designed to effect a bio-
logical control of algae and, specifically, to
determine the basis for the resistance and
susceptibility of algae to microbial attack.
We thank I. Feigenoff for her assistance.
Support for the work was provided by a
Ford Foundation training grant in the ecol-
ogy of pest management.
Methods and materials
The algae, all in axenic culture, were
selected to represent a range of planktonic
and attached species, but the choice was
also governed by the ability of the organ-
isms to grow in the fermentors used and
their suitability for harvesting by continu-
ous flow centrifugation, Algae were grown
in 250 ml of a modified Bristol’s medium
(BBM) (Bold 1942) in 500-ml Erlenmeyer
flasks at 22°C on a rotary shaker at 150
l Present address: Waterways Experiment
tion, U.S. Army Corps of Engineers, Vicksburg,
Mississippi 39180.
LIMNOLOGY AND OCEANOGRAPHY
rpm under 5,400 lux constant illumination
for 18 to 23 days and used to inoculate an
18-liter carboy containing 16 liters of BBM.
The culture was incubated at 22°C with
illumination at 10,800 lux provided by two
fluorescent lamps mounted 5 cm from the
sides of the bottle. Filter-sterilized air was
passed through the medium at a rate of 10
liters min-l. The cultures were harvested
after 19 to 23 days by centrifugation, and
the cells were washed three times with cold
0.01 M phosphate buffer, pH 7.0, and then
suspended in 100 ml of the buffer. The
final suspension was termed the “washed
algal concentrate” and was either used
immediately or lyophilized and stored at
-17°C.
To test the susceptibility of algae to
decomposition, we used a method modi-
fied from that of Gromov and Mamkaeva
(1969). Washed algal concentrate (2.5 ml),
tither living or steamed for 20 min, was
spread evenly on the surface of BBM solidi-
fied with 2.0% agar in a petri dish to form
a “lawn.” The preparation was then per-
mittcd to dry for 4-5 days at 20°C under
10,800 lux illumination. Samples of soil and
sediment (0.1 g) or natural waters (0.1 ml)
were added to the surface of the algal
1awns, with sterilized samples serving as
Sta-
controls. The soil was either Williamson
silt loam from a cultivated field or a garden
64 JANUARY 1975, V. 20( 1)

Microbial decomposition
soil. Sediment samples came from a fresh-
water swamp or a water-filled ditch. Water
was collected from Cayuga Lake, Jenning’s
Pond, or the primary settling tank of a
sewage treatment plant. The plates were
incubated in a moist atmosphere at 30°C
in the dark for 30 days and examined daily
for evidence of discoloration, visual disap-
pearance of algal biomass, loss of algal
walls, and release of algal cell contents.
To dcterminc the validity of the labo-
ratory findings, WC conducted a study in
Cornell University experimental pond 201.
This pond has a surface area of about 0.07
ha and a mean depth of 1.5 m (Hall et al.
1970). The pond is mesotrophic, lacks any
emergent vegetation except Ear a stand of
Typha sp. in one corner, and has Chara sp.
as the dominant bottom plant.
A covered raft was placed in the middle
of the pond which held 18 removable sam-
ple blocks submerged beneath the surface.
Affixed vertically to each side of 15 of the
blocks were 14 Pastcur pipets, with the
wide end downward and the tips removed
to provide openings of 2-3 mm. The wide
ends were packed with glass wool to a
tightness sufficient to prevent the passage
of algae while permitting the entry of wa-
ter and small microorganisms. Into each
pipet was introduced 1.0 ml of washed al-
gal concentrate; the glass wool plugs re-
tained the algae while the buffer drained
out. All 14 algae were on each side of the
block, and each alga was thus represented
twice on one block. The pipcts were kept
in darkness about 10 cm beneath the pond
surface. The aquatic microbiota had access
to the algae through the unobstructed top
of the pipct, and the glass wool plug at the
bottom was sufficiently loose to permit the
exchange of nutrients and dissolved gases
from the water. Controls were provided
by lo-ml ampoules containing 2.0 ml of
washed algal concentrate and 6.0 ml of
filter-sterilized pond water. The ampoulc
tops were sealed with sterile parafilm, and
the ampoules were fastened to the rcmain-
ing 3 sample blocks and held beneath the
water surface at the same depth as the
pipcts.
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of algae 65
A sample block of 28 pipets was removed
from the raft every 7 days for 12 weeks,
and the algal layer was examined micro-
scopically. At the same time about 0.5 ml
of the control suspension was removed from
each ampoule aseptically for examination,
but the control ampoules were then re-
turned to the raft.
To test the ability of culture filtrates to
inhibit microbial growth, we sterilized the
cell-free liquid from a 21-day algal culture
by filtration and introduced portions of this
liquid or sterile 13BM into stainless steel
antibiotic assay cylinders supported on an
agar medium previously seeded with a test
heterotroph. The plates were incubated
for 2 to 5 days at 30°C and examined for
the appearance of clear zones around the
cylinders.
The existence of intracellular compounds
with antimicrobial activity was also investi-
gated. The washed algal concentrate was
suspended in an equal volume of distilled
water and disrupted by treatment for 15
min with a sonic oscillator at a temperature
not exceeding 20°C. The resulting suspen-
sions or those remaining after centrifuga-
tion at 3,000 x g were placed in antibiotic
cylinders and assayed as above.
The method of preparing the cell walls
of Fischerella muscicola, Pediastrum du-
plex, Staurastrum sp., and [Jlothrix fim-
hrata, cell envelopes of Cylindrospermum
SP*, and wall fragments of Chlamydomonas
reinhardtii will be described elsewhere
( Gunnison and Alexander in prep. ) . The
procedure involved repeated rupture of
the cells with a sonic oscillator (or a French
pressure cell for C. reinhardtii) followed by
extensive washing of the material collected
by centrifugation; the wall preparations
were then suspended in sodium dodecyl
sulfate, treated again by sonic oscillation,
and finally suspcndcd in distilled water.
For the isolation of microorganisms able
to degrade cell walls, the salts solution of
Potgieter and Alexander (1966) was sup-
plemented with 0.001% yeast extract and
0.1% algal ccl1 walls, 0.2% lyophilized algal
cells, or 0.2% of either xylan or MN cellu-
lose 300 (Brinkmann Instruments). The
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66 Gunnison and Alexander

Table 1. Susceptibility of lawns of live algae to microbial decomposition.

Days for decomposition to be evident

Source Soils Sediments Waters
Al gafi of alga.1 Field Garden Swamp Ditch Lake Pond Sewage

Anabaena flos-aquae (Cya) 3 3 4 4

ms nidulans
Ankistrodemcatus
(Cya)
(Chl )
IU
IU 1 2 2 i6 28
Calothrix
Chlamydomonas
anomaw)
reinhardtii (C hl )
IU
CB
CU
10
>30
6
10
>30
4
1;
3
11
25
9
>30
6
330
6
12
8
Chlorella sp. (Chl) CU 17 16 20 24 '30 >30 9
Cylindrospermum sp. (Cya) 5 7 10 10 7 6
Euglena
Fischerella
gracilis (Eug)
muscicola
IU
IU
CB
11
>30
9
>30
14
>30
11
>30
13 ii 5
>30 >30 >30
IU
IU
CAMB >30
:
>30
;
730
i ::
>30 730
2 12
16
'30
6
6
>30
IU 20 24 22 22 27 26 9
--Ulothrix fimbrata (Chl) CAMB 10 9 10 13 6 5 4

%hl: Chlorophyta; Cya: Cyanophyta; Eug: Euglenophyta.

TCAMB: Cambridge Univ. Culture Collection of Algae and Protozoa; CB: Carolina Biological Supply
co. ; CU: Cornell Univ. collection; IU: Culture Collection of Algae at Indiana Univ.

solution was inoculated with a sample of
soil or aquatic sediment and incubated at
30°C. Serial transfers were made to fresh
medium when much of the wall material
had disappeared or after 40 days, which-
ever came first; after several such transfers,
the enrichment was streaked onto the salts
medium solidified with 2.0% agar and over-
lain with a suspension of 0.1% walls or
0.2% algal cells in 1.5% agar. The plates
were incubated for l-5 days, and colonies
surrounded by clear zones were picked and
identified by the methods described by
Skerman ( 1967).
Results
Decomposition in algal lawns
and natural water
Viable algae present on an agar surface
differed markedly in their susceptibility to
degradation by microorganisms present in
soils, sediments, and bodies of water (Ta-
ble 1). The values given represent the
times elapsed before decomposition was
evident microscopically; each is the aver-
age of three replicates. Observations ended
after 30 days. The data show appreciable
differences among the algae in their resis-
tance to digestion by microorganisms, the
cells of some being rapidly destroyed while
others are quite refractory to attack. Dif-
ferences are evident also in the rates of di-
gcstion of a single alga by organisms from
the different habitats, but no consistent
trend is evident.
Anabaena flos-aquae and Anacystis ni-
dulans were quite susceptible to microbial
decomposition, the average time for their
digestion, considering samples from all
the environments, being less than 5 days.
Ankistrocbsmus falcatus, C. reinhardtii,
Cylindrospermum sp., Eugleena gracilis,
Gloeocystis &as, Ourococcus multisporus,
and U. fimbrata were somewhat more re-
sistant; the average time for the initiation
of their decomposition ranged from 6-10
days. Stazwastrum sp. was less liable to
degradation with generally more than 20
days before decomposition was observed.
Calothrix anomala and Chlorella sp. were
likewise moderately resistant, and attack
was occasionally not noted at all in the test
period. On the other hand, I;. muscicola
and P. duplex were never attacked appre-
ciably during the study period.
The patterns of resistance and suscepti-
bility were quite similar when dead algae
were used (Table 2). The values again
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Microbial decomposition of algae 67

Table 2. Susceptibility of lawns of killed algae to microbial decomposition.

Days for decomposition to become evident

Soils Sediments Waters
Alga Field Garden Swamp Ditch Lake Pond Sewage

4
Anabaena
ms
flos-aquae

Ankistrodesmus
nidulans
falcatus
3
5

:6 : 2
8
3
3
7
Calothrix anomala 7 :: 1; 15 i
Chlamydomomnhardt -ii 8 7 8 6 4
Chlorella
Cylindrospermum
sp. -
sp.
197
6
17 13
5
16 24 28
8
4
Euglena gracilis
Fischerella muscicola
11
r30 730
z . 19
730
95
>30
1;
730
11
r30
:
23
Gloeocystis CJ&& 12 11 13 10 8 9 4
Ourococcus multisporus 5 5 5 5 8 8 4
Pediastrum duplex >30 >30 >30 r30 >30 >30 p30
Staurastrum sp. 230 >30 >30 22 26 >30 15
--Ulothrix fimbrata 9 6 9 13 8 7 3

represent the averages of three replicates.
Again taking into account the times for di-
gestion for samples from all environments,
A. flos-aquae and A. nidulans are most
prone to digestion. As with the viable al-
gac, the average times for evidence of an
attack 011A. falcatus, C. reinhardtii, Cylin-
drospermum sp., E. gracilis, G. gigas, 0.
multisporus, and U. fimbrata were 6-10
d ays. Nonviable cells of Chlorella sp. were
still less susceptible, but the resistance of
C. anomala was markedly reduced when
the cells were steamed. Fischerella musci-
cola, P. duplex, and Staurastrum sp. were
resistant whether viable or heated; digcs-
tion of P. dupZex was never observed. The
cells of algae such as A. falcntus and Chlo-
rella sp. also generally lost some of their
resistance as a result of steaming, whereas
the resistance of G. gigas and Staurastrum
sp. usually increased as a conscqucncc of
this treatment.
Marked differences in resistance to de-
composition were evident in the pond wa-
ter (Table 3); many spccics were dcstroycd
with comparative ease, while others per-
sisted for 2-3 weeks. Staurastrum sp, and
F. muscicola showed no signs of signifi-
cant attack until the fourth week, and P.
duplex endured for 12 weeks, when the
study ended.
For comparative purposes, the ranges of
values for the lawns of viable algae arc also
prescntcd in Table 3. The data for the two
means of testing for degradability are in
general agreement, but a few disparities
exist. For example, C. anomala was not
subject to discernible attack for the first 18
days on lawns but was readily decomposed
in the natural pond. Decomposition of P.
duplex was never found, either in the 30-
day laboratory trial or in the 12-week field
evaluation.
Possible rcsis tancc mechanisms
As an explanation for the resistance of
some of the algae, it sccmcd plausible that
some spccics might excrete extracellular
toxins that prevent the growth of lytic, par-
asitic, or predatory organisms. I-Iowcver,
culture filtrates had no discernible effect
on the development of Aspergillus niger,
Penicillium brefeldianum, or Saccharomy-
ces cerevisiae on potato dextrose agar, Ba-
cillus subtilis, Micrococcus lysodeikticus,
Nocardia sp., or Streptomyces sp. on nutri-
ent agar, or Rhixobium sp. on mannitol-
yeast extract agar. These heterotrophs were
selected to reprcscnt a diverse group of
organisms so that antimicrobial activities,
if prcscnt, might bc observed.
Diffcrcnccs in susceptibility to attack
might also result from the presence of in-
tracellular algal constituents inhibitory to
potential lytic, parasitic, or predatory or-
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68 Gunnison and Alexander

Table 3. A comparison of the susceptibility of inoculation. Daily examination of these

live algae to decomposition in a pond and in inoc-
plates failed to reveal any depression in
ulated lawns.
-.
the growth of the individual heterotrophs
Days for the initiation
or of members of the soil or sediment
of decomposition communities.
Alga Pond;: Algal lawns? To determine the effectiveness of the cell
wall as a barrier to agents of biodegrada-
tion, WC tested the suitability of algal wall
Anabaena
ms
Ankistrodesmus
flos-aquae
nidulans
falcatus 14
2
6-19
preparations as a substrate for microorgan-
Calothrix anomala 19-730 isms. Uecause of the number of steps in-
Chlamydomoxnhardti i 2; 3-9 volved in obtaining an uncontaminated wall
Chlorella sp. g-730
Cylindrospermum sp. preparation, only six algae were used: C.
Euglena gracilis li ::1:: reinhardtii, Cylindrospermum sp., and U.
Escherella muscicola 28 730
Gloeocystis gigas 14 4- 12 fimbrata were selected as susceptible algae,
Ourococcus multisporus 7 3-16 Staurastrum sp. as a moderately resistant
Pediastrum duplex ~84 930
Staurastrum sp. 26 9-27
organism, and I?‘. muscicoZa and P. duplex
Ulothrix
-- fimbrata 7 4-13 as the more refractory species. When walls
of the susceptible algae were introduced
into enrichment cultures, they underwent
+:Sampling performed at weekly intervals.
?The range of times for degradation to be detect- some degree of decomposition. The walls
able. of C. reinhardtii and Cylinclrospermum sp.
disappeared completely within 12 days.
Walls of U. fimbrata dccomposcd somc-
ganisms. However, no unfractionated prcp- what more slowly, but after 28 days of in-
arations made by treating the algae with cubation looked ragged and perforated and
a sonic oscillator and none of the same had become noticeably thinner than they
preparations subjected to centrifugation at were originally. The walls of Staurastrum
3,000 x g inhibited the growth on agar of sp., F. muscicola, and P. duplex did not
the heterotrophs listed above. In addition, appear to bc attacked, even after 40 days
extracts of cells of the resistant algae wcrc in the cnrichmcnts.
mixed with an equal volume of nutrient
broth prepared with twice the normal
Digestion of walls
concentration of ingrcdicnts, and this mix-
ture was inoculated with samples of garden Eight different microorganisms capable
soil, field soil, swamp sediment, or synthetic oE digesting walls of the susceptible al-
sewage prepared by James’ (1964) method, gae were obtained from soil. The carbon
inoculated with a natural sewage micro- sources used for the enrichments were C.
flora, and incubated at 30°C for periods reinhardtii walls for LMicrobacterium sp.;
up to 5 days. Here too heterotrophic C. reinharcltii cells for a Corynebacterium
growth was not detectably affected by the and Brevibacterium G2; xylan for Brevi-
algal constituents. bacterium G5; and C ylintlrospermum sp.
In another test of the possible formation walls, TI. fimbrata walls, and ccllulosc for
by the algae of antimicrobial compounds, Streptomyces strains G6 and F7, G4, and
the heterotrophs listed above or samples D8, respectively. Grcatcr difficulty was
of soil or sediments were inoculated onto encountered in finding isolates able to uti-
lawns of the various algae. The agar me- lize U. fim7jrata walls than in obtaining
dium was prepared by placing 2.0 ml of heterotrophs using the surface layers of
a mixture containing equal volumes 0E a the other susceptible algae, and the cleared
thick algal suspension and tenfold conccn- zones produced by the bacterial and acti-
tratcd nutrient broth on top of 10 ml of nomyce tc colonies growing on plates with
13RM agar in a petri dish and allowing the [J. fim7jrata walls invariably contained rc-
plates to dry for 2-5 days at 20°C before sidual wall components, although some
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Microbial decomposition of algae 69

Table 4. Digestion of algal walls by individual microbial rcsidcnts of pond water. Fisch-
microorganisms. ereZZamuscicola was the only resistant blue-
green of the five examined, but still it was
C. rein-
Digestion of walls
Cylindro-
of
i. fim-
attacked to some extent after 4 weeks in
Isolate rardtii spermum sp. brata natural waters. The unsuitability of the
blue-green algae as food for higher organ-
Microbacterium sp.
+;" -I- isms is often cited, in particular for those
+;':
Brevibacterium
Corynebacterium
G2
sp.
+"' planktonic crustaceans grazing on algae
Streptomyces D8
+A
+ + (Hutchinson 1973). The grazing pressure
+': -E
Streptomyces G6
+
+:"
-4
cxcrtcd by zooplankton on blue-greens
Streptomyces F7
Brevibacterium G5 -I- + +':' ranged from variable to ineffective (Porter
+ + +;;
Streptomyces 64 1973 ) ) although a few blue-greens are
by themsclvcs able to meet the nutritional
;:Walls serving as carbon source for original isola- requirements of Daphnia pulex (Arnold
tion. 1971). I3y contrast, blue-green algae are
known to bc consumed, lysed, or parasit-
degradation and thinning of the surface ized by a wide range of microorganisms.
components were always evident. We For example, Ho and Alexander (1974)
foulld two microorganisms which caused reported the utilization of blue-greens as
this kind of digestion of 77. fimbrata walls. food sources by several spccics of amcbac.
We found no isolates that could grow on Viruses acting on blue-greens have been
or dissolve the walls of any of the three foulid ( Shilo 1971)) and bacteria lysing
resistant algae. blue-green algae have been isolated fre-
The bacteria and actinomycetcs wcrc quently (Daft and Stcwart 1971; Granhall
tested for their ability to digest algal walls and 13crg 1972; Gromov et al. 1972; Shilo
other than those present in the mediuln for 1970). TIcnce it is not mlexpectcd that
their original istilation. The isolates were most of the blue-greens we tested are sus-
streaked on a plate containing an inorganic ccp tiblc to decomposition.
salts-agar medium on top of which was Among the Chlorophyceac examined,
placed water agar supplemented with the only C. reinharcltii and 0. multisporus wcrc
cell walls. Walls of none of the resistant particularly susceptible to decomposition,
algae were solubilized or digested to an cx- and the green algae we studied wcrc gen-
tent evident as zones of clearing around the erally modcratcly or highly resistant. The
streaks. 13~ contrast, four of the eight iso- resistance of these Chlorophyceae might
lates degraded walls of all the susceptible seem to contradict the fact that many of
algae (Table 4), although the walls used these algae serve as food for grazing or-
for the initial isolation were always the ganisms. IIowever, those algae that fall in
more cxtcnsivcly destroyed. Two digested the susccptiblc to susceptible-intermediate
only the walls on which they were initially category in our studies arc often subject to
isolated. heavy grazing pressure. Chlamydomonas is
digested by zooplankton (Porter 1973); I-IO
Discussion and Alexander ( 1974) found that C. rein-
The rcsistancc of algae to microbial de- hardtii, 0. multisporus, and A. falcatus
composition may well be diffcrcnt from were consumed by various amebae, a1-
their suitability as food sources to higher though Chlorella sp. and Staurastrum sp.
organisms. For example, our results show were complctcly resistant to the protozoa.
that three of the genera of blue-green algae Arnold ( 1971) found that both A. falcatus
arc particularly susceptible to microbial at- and Chlorella vulgaris were capable of sup-
tack under all conditions tried. Even C. porting D. pulex; however, larger green
anomala, although somcwhat slowly uti- algae and desmids (like Staurastrum) arc
lized on lawns, was readily decomposed by ullaffccted by grazing pressure, lullike the
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70 Gunnison and Alexander

smaller nongelatinous green algae, which BOLD, H. G. 1942. The cultivation of algae.
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DAFT, M. J., AND W. D. P. STEWART. 1971.
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and it is quite possible that algal resistance a1gac. New Phytol. 70: 819-829.
is associated with toxic metabolites. The GLOMHTZA, K.-W, 1970. Antimikrobiclle In-
toxins can affect other algae ( Lefevrc et haltsstoffc in Algen. 2. Das Vorkommcn van
al. 1952) or nearby heterotrophs (Pratt Acryls#ure in verschicdenen Mcercsalgcn.
Planta Med. 18: 210-221.
et al. 1944). The toxicity may sometimes GI~ANIIALL, U., AND B. BERG. 1972. Antimicro-
even be attributable to the high oxidation- bial effects of CeZZ&brio on blue-green algae.
reduction potentials resulting from algal Arch. Mikrobiol, 84<: 234-242.
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Fcdenko 1965). The results of our study AND I. A. AVILOV. 1972. Flexibacterium lys-
ing blue-green algae. Mikrobiologiya 41:
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intracellular, or localized at the cell sur- - AND K. A. MAMKAEVA. 1969. Sensitivity
face-are not important in regulating the of ‘diffcrcnt Scenedesmus strains to the endo-
growth of potential microbial agents of dc- parasitic microorganism Amoebo~$eZicZiz~m.
composition, at least for the algae we inves- Phycologia 7 : 19-23.
tigated. In nature, nevertheless, toxins may IIALL, D. J,, W. E. COOPER, AND E. E. WERNER.
1970. An experimental approach to the pro-
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Our results do suggest, however, that the
10 : 95-100.
cell wall is a major determinant of algal I~UTCIIINSON, G. E. 1973. Eutrophication. Am.
resistance. Although walls of the readily Sci. 61: 269-279.
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Submitted: 11 March 1974
Oceanogr. 16 : 906-920. Accepted: 22 August 1974