1. Title of paper and include the link if it is one you chose versus one I assigned
2. List 3 observations from the paper that inspired the authors to conduct the experiment or write this paper. DO NOT tell me what they did. Tell me WHY they are even going the study – 3 reasons that should be based on previous research and is usually addressed in introductions.
3. What was/is their over-arching question?
4. What is their hypothesis?
5. What was their independent variable?
6. What was their dependent variable?
7. EXPLAIN what they did in their experiment in your own words. This will require some details, show me you understood everything they did.
8. What was their control, did they have one?
9. Summarize the results of their data.
10.
Summarize why this study matters? What does it tell us? Why should we care?
O R I G I N A L P A P E R
Competitive exclusion of Cyanobacterial species
in the Great Salt Lake
Hillary C. Roney Æ Gary M. Booth Æ
Paul Alan Cox
Received: 25 July 2008 / Accepted: 16 December 2008 / Published online: 8 January 2009
� Springer 2009
Abstract The Great Salt Lake is separated into different
salinity regimes by rail and vehicular causeways. Cyano-
bacterial distributions map salinity, with Aphanothece
halophytica proliferating in the highly saline northern arm
(27% saline), while Nodularia spumigena occurs in the less
saline south (6–10%). We sought to test if cyanobacterial
species abundant in the north are competitively excluded
from the south, and if southern species are excluded by the
high salinity of the north. Autoclaved samples from the
north and south sides of each causeway were inoculated
with water from each area. Aphanothece, Oscillatoria,
Phormidium, and Nodularia were identified in the culture
flasks using comparative differential interference contrast,
fluorescence, and scanning electron microscopy. Aphanot-
hece halophytica occurred in all inocula, but is suppressed
in the presence of Nodularia spumigena. N. spumigena was
found only in inocula from the less saline waters in the
south, and apparently cannot survive the extremely
hypersaline waters of the northern arm. These data suggest
that both biotic and abiotic factors influence cyanobacterial
distributions in the
Great Salt Lake.
Keywords Competitive exclusion � Halophilic bacteria �
Aphanothece � Nodularia � Oscillatoria � Phormidium �
Gause’s principle
Introduction
Cyanobacteria are well-adapted for living in harsh condi-
tions, including photosynthetic areas beneath Antarctic ice,
hot springs and geysers in Yellowstone, and hypersaline
lakes, including the Great Salt Lake (Dyer 2003). Two
common cyanobacteria species that have been identified
from the Great Salt Lake are Aphanothece halophytica and
Nodularia spumigena (Brock 1976; Felix 1978; Felix and
Rushforth 1980), with N. spumigena episodically blooming
in Farmington Bay (Marcarelli et al. 2006). Since nitrogen
is the limiting nutrient in the Great Salt Lake (Oren 2002) it
is interesting to note that both A. halophytica and
N. spumigena can fix nitrogen.
The Great Salt Lake is a hypersaline remnant of the
Pleistocene Lake Bonneville (Oren 2002) which was
557 km long and 233 km wide with an area of 51,800 km
2
in what is now Utah, Idaho, and Nevada (Utah Geological
Survey 1990). As Lake Bonneville retreated, the lake lost
all outlets, so salinity increased.
After completion in the mid 19th century of the
transcontinental railway near Promontory Point, Utah,
trains had to traverse many additional rail kilometers
around the northern end of the Great Salt Lake. To reduce
this distance, the Union Pacific Railroad constructed a
19 km rail causeway across the Great Salt Lake in 1959
(Fig. 1), replacing an earlier trestle built in 1902 by the
Southern Pacific Railroad. Unlike the former wooden
trestle, which did not impact water flow, the 1959
causeway built with rock fill, hydrologically divided the
Communicated by L. Huang.
H. C. Roney � P. A. Cox (&)
Institute for Ethnomedicine, Box 3464,
Jackson Hole, WY 83001, USA
e-mail: paul@ethnomedicine.org
G. M. Booth
Department of Plant and Wildlife Science,
Brigham Young University, Provo, UT 84602, USA
123
Extremophiles (2009) 13:355–361
DOI 10.1007/s00792-008-0223-1
waters of the Great Salt Lake into two portions, a northern
arm with negligible freshwater inputs, and a southern arm
with more than 90% of the freshwater flow (Butts 1980;
Oren 2000; Stephens and Gillespie 1976; Sturm 1980).
Three major rivers–the Bear, Weber, and Jordan–all flow
into the Great Salt Lake south of the railway causeway
(Gwynn 2002). The overall salinity of the Great Salt
Lake, which to that point had been linked solely to
changing water levels, quickly adjusted with the northern
arm becoming even more hypersaline, moving from an
average of 15% in the 1870s to as high as 28% salinity in
the 1960s (Sturm 1980). In 1970, the northern arm held
approximately 330–350 g salts per liter, while the south
arm held 120–130 g salts per liter (Oren 2002). While the
major cation in the water is Na, Mg, K, Ca, in decreasing
order of abundance are important as is the anion SO4
(Sturm 1980).
These habitat changes were later partially replicated
with construction of a second barrier to lake water flow.
A causeway for vehicular use has been periodically
constructed from Syracuse, Utah to Antelope Island, and
was most recently rebuilt in 1992 (Gwynn 2002). These
vehicular and railway causeways resulted in the parti-
tioning of the Great Salt Lake into three different salinity
regimes: the northern arm, with an average of 27%
salinity, the middle arm with average salinity of 10–16%,
and the southern arm, with average salinity of 6% or less
(Utah Geological Survey 1990). These three different
salinity regimes, any one of which would be considered
hypersaline, allowed species to sort according to eco-
logical tolerances. The results are striking: each large
area of the lake has different colored water, resulting in
part from different concentrations of Artemia fransiscana
brine shrimp cysts and microscopic green algae such as
Dunaliella salina and D. viridis as well as species of the
Archean genus Halobacterium (Post 1981), but also
perhaps due to different concentrations and species
compositions of cyanobacteria.
A study was designed to determine whether cyanobac-
terial distributions in the Great Salt are influenced by
abiotic factors, biotic factors, or both. To explore this
question, experiments were designed to examine two
hypotheses; Hypothesis 1: Cyanobacterial species abundant
north of the railway causeway are competitively excluded
from the south by other species, and Hypothesis 2:
Cyanobacterial species that thrive and bloom south of the
Antelope causeway cannot grow in high salinity waters
from the north.
Materials and methods
Experimental cultures
A total of 28 water samples from both the north and south
sides of Antelope Island causeway and the north and south
sides of the railway causeway (seven from each site) were
collected in December 2007. Water temperatures and GPS
coordinates were recorded at each site. To avoid pseu-
doreplication, six water samples from both sides of the
vehicular and railway causeways, approximately four liters
in volume, were used as inocula. The seventh jar from each
side was approximately eight liters in volume and used for
media after filtering and autoclaving. Approximately 30 ml
of the filtered media water was placed in autoclavable,
sterile 50 ml nalgene plastic flasks. In total there were 96
flasks inoculated with 10 ml of unsterilized water from
either the north or the south of the railway and the vehic-
ular causeways totaling six replicates of inoculum water for
each medium type. No nutrients or other growth media
were added to the water (Dyer 2003). In addition, 21
control flasks were prepared using autoclaved media and
autoclaved inocula. A random number table was used to
decide from which sample jars to draw the inoculum. In
addition, one control flask was prepared which consisted of
autoclaved distilled water inoculated into autoclaved dis-
tilled water medium. After all 129 flasks were inoculated;
they were placed in a heated green house with constant
8.5 h/day illumination. Each flask was gently shaken by
hand periodically for aeration. These liquid cultures were
incubated for 7 weeks.
Cyanobacterial identification
For aquatic cyanobacteria, identification by light micros-
copy (phase and/or interference contrast), and scanning
electron microscopy (SEM) are preferred (Cronberg and
Fig. 1 Great Salt Lake Rail Causeway with hypersaline water in the
north (left) and less saline water on the south (right)
356 Extremophiles (2009) 13:355–361
123
Annadotter 2006). Identification and abundance counts of
cyanobacteria from the culture flasks were performed using
differential interference contrast (DIC) and epi-fluores-
cence imaging, with SEM for verification.
Data analysis
To ensure arbitrariness of cyanobacterial counts, micro-
transects of water cultured from each flask, based on two
microscope slides were conducted. Each microtransect
was replicated twice for each slide, with data entered on a
six-cell mechanical lab counter. When mass colonies of
cyanobacteria where encountered precluding individual
counts, the colony were assessed as ‘‘large’’ or ‘‘very
large’’, with medians and nonparametric analyses used to
analyze qualitative data. In each of the 16 possible
combinations of 4 types of media (railway north, railway
south, Antelope north, and Antelope south) and 4 types of
inocula (railway north, railway south, Antelope north, and
Antelope south), the median counts of the 4 major
cyanobacterial species (A. halophytica, Oscillatoria sp.,
Phormidium tenue, and N. spumigena) were ranked.
A two-way Analysis of Variance (ANOVA) was cal-
culated for abundance of A. halophytica in the culture
flasks with ‘‘large’’ colonies scored as 500 and ‘‘very
large’’ colonies scored as 1,000 for this purpose. To reduce
impact about outliers and ensure consistency of distribution
across the observed range, all data were transformed with a
square root transformation prior to analysis as is standard
for count data. F statistics for the transformed data were
calculated to test three different pairs of hypotheses, with
the null hypothesis to be rejected at the P \ 0.05 level:
Hypothesis pair #1
H0: no variation in cyanobacterial counts exists due to
differences in media.
H1: variation in cyanobacterial counts exists due to
differences in media.
Hypothesis pair #2
H0: no variation in cyanobacterial counts exists due to
differences in inocula.
H1: variation in cyanobacterial counts exists due to
differences in inocula.
Hypothesis pair #3
H0: no variation in cyanobacterial counts exists due to
interactions.
H1: variation in cyanobacterial counts exists due to
media and inocula interactions.
For cyanobacterial taxa which proved to be of rare
occurrence in the culture flasks, exact logistic tests, rather
than an ANOVA, were calculated.
Results
Experimental cultures
When sampling for the experimental cultures of the Great
Salt Lake, profoundly different colors on either side of the
railway causeway were observed from the air (Fig. 1).
These differences were also apparent in water samples
taken from deep water on either side of the causeway rather
than evaporative ponds (Fig. 2). Salinity from the sample
sites was previously measured by hydrometers—south side
of Antelope causeway 20.6 ppt, north side of Antelope
causeway 75 ppt, south side of railway causeway 155 ppt,
and north side of railway causeway 195 ppt (Roney 2007).
Salinity values of the flasks were altered slightly by addi-
tion of inocula, except when the same inoculum was added
to the same medium.
At the time of sampling in December 2007, ambient air
temperature was -2.2�C on the railway causeway
(4181301600N1128320303400W) and -2.8�C on the vehicular
causeway (418404400N11281205700W), with water tempera-
tures north of the railway causeway at 2.0�C, south of the
railway causeway at 4.3�C. Water temperature north of the
vehicular causeway was 3.3�C, while the water tempera-
ture south of the causeway was 2.5�C. Analysis by light
microscopy showed no growth in any of the 21 control
flasks, which were found to be sterile.
Cyanobacterial identification
Four genera of cyanobacteria, Aphanothece, Oscillatoria,
Phormidium and Nodularia, were identified (Figs. 3, 4, 5,
6), with identifications confirmed by Dr. James Metcalf
Fig. 2 Water samples taken on north (left) and south (right) side of
railway causeway. Color differences primarily due to Dunaliella
distributions although the cyanobacterium Aphanothece flourishes in
the hypersaline waters in the north
Extremophiles (2009) 13:355–361 357
123
(University of Dundee, Scotland). In addition, a fifth
cyanobacterial genus, Spirulina, was observed, but was not
found in any transect through any of the microscope slides.
Because of its trichomes and its affinity for saline waters,
this species is referable to Spirulina labyrinthiformis
(Fig. 7), although Nübel et al. (2000) have placed a similar
salt-tolerant species into the new genus, Halospirulina.
Comparisons between differential interference contrast
microscopy, fluorescence microscopy, and scanning elec-
tron microscopy allowed different observations of
cyanobacterial morphology and size to be compared for
taxonomic identification.
Data analysis
Comparative medians of the four cyanobacterial genera for
each inocula type in the four media are shown in Fig. 8,
which demonstrates that A. halophytica appears throughout
all the four types of media and inocula, but that Nodularia
spumigena is abundant only in inocula from Antelope south
waters. Since there are 24 different permutations of the
ordered ranks of Aphanothece (A), Oscillatoria (O), Nod-
ularia (N), and Phormidium (P) plus an additional 16
permutations of three and two-way ties, as well as one
Fig. 3 Aphanothece halophytica: differential interference contrast
(left); fluorescence (middle); scanning electron microscopy (right)
Fig. 4 Oscillatoria sp.: differential interference contrast (left); fluo-
rescence (middle); scanning electron microscopy (right)
Fig. 5 Phormidium tenue: differential interference contrast (left);
fluorescence (middle); scanning electron microscopy (right)
Fig. 6 Nodularia spumigena: differential interference contrast (left);
fluorescence (middle); scanning electron microscopy (right)
Fig. 7 Spirulina cf. labyrinthiformis: differential interference con-
trast (left); fluorescence (middle); scanning electron microscopy
(right)
358 Extremophiles (2009) 13:355–361
123
possible case of a four-way tie in rank, there are 41 dif-
ferent possible rankings of the four cyanobacterial species.
These different rankings can perhaps most easily be por-
trayed as different colors (Fig. 8). A. halophytica was
dominant in all cultures flasks, except those in which N.
spumigena and Oscillatoria sp. occurred.
A two-way ANOVA for the distributions of A. halo-
phytica was performed as indicated in Table 1. The F
statistics for the ANOVA allows each of the null hypoth-
eses in the three pairs of hypotheses to be rejected at the
P \ 0.05 level. Therefore, it can be concluded that media
and inocula, as well as the interaction between media and
inocula significantly affected the growth of A. halophytica
in the culture flasks. Exact tests were calculated for
N. spumigena and Oscillatoria spp. using the exact option
of pro logistic in SAS. In these analyses, counts were
ignored, and instead, presence/absence data were used. For
Oscillatoria, the P value for the exact test of the medium
was 0.0993; thus the effect due to media differences was
not significant. However, the P value for the exact test of
inoculum was 0.0016; hence there was a significant inoc-
ulum effect in distribution of Oscillatoria. The exact test
for the inocula/media interactions was not significant with
a P value of 0.1963. Leaving interaction out of the model,
the additive model (with additive effects of inoculum and
medium) showed the odds of a positive response for
inocula from rail south, rail north, or Antelope north was
just 6.3% relative to inoculum from Antelope south (95%
confidence interval: 0.6–64.2%). Thus, Antelope south
inocula had a significant positive effect on the presence of
Oscillatoria in the culture flasks.
A similar analysis was conducted for the presence or
absence of N. spumigena in the culture flasks. The exact
test for the inocula/media interaction had a P value of
0.0032; hence the interaction was significant. The odds of a
positive effect were extremely high for the combination of
Antelope south inoculum with Antelope south medium. For
all other combinations, the odds of a positive response were
extremely low. For future studies of algal-cyanobacterial
interactions, counts were also made of the green alga
Dunaliella salina and D. virids in the flasks; an ANOVA of
square root transformed data count for Dunaliella showed
significant differences in distributions similar to Aphanothece
distributions; these data will be reported elsewhere.
Discussion
Both the ranking of median abundances in the culture
flasks and the results of the two-way ANOVA support the
overall hypothesis: cyanobacterial species abundant north
of the railway causeway (e.g. A. halophytica) are com-
petitively excluded from the south by other species, in this
case N. spumigena and Oscillatoria spp. It appears that the
cyanobacterium A. halophytica can grow in less saline
waters as well as the extreme saline waters north of the
railway causeway—since it is found in all inocula—but its
growth appears to be suppressed in the south by the pres-
ence of N. spumigena, which periodically blooms in the
Great Salt Lake.
In previous years, we have noted large N. spumigena
blooms in the low salinity regime of Farmington Bay, as well
as in water samples collected south of the railway causeway
(Roney 2007), particularly when winds have concentrated
blooms near the causeway. Rushforth and Felix (1982)
recorded N. spumigena as rare in the south arm; perhaps they
took their samples at a dormant season, as N. spumigena
blooms episodically. The absence of N. spumigena in the
southern arm of the Great Sale Lake may have influenced
the ability of A. halophytica to migrate and prosper in the
fresher water environment of the south arm instead of
thriving in the hyper-saline north arm. In all of our samples
south of the Antelope causeway (Farmington) since 2004,
N. spumigena was present in the water column.
The second overall hypothesis—that cyanobacterial
species that thrive and bloom south of the Antelope
Fig. 8 Rankings of cyanobacterial dominance by medians. Media/
Inocula in the upper left corner of the chart are extremely hypersaline,
while those in the lower right corner are far less saline
Table 1 ANOVA of Aphanothece halophytica distributions in
autoclaved media from the Great Salt Lake
Source Variation Degrees
free
Mean
square
F
Statistic
Significance
Media 64.6 3.0 21.5 11.4 P \ 0.01
Inocula 26.1 3.0 8.7 4.6 P \ 0.01
Interaction 105.6 9.0 11.7 6.2 P \ 0.01
Subtotal 196.3 15.0
Error 151.1 80.0 1.9
Total 347.4 95.0
Extremophiles (2009) 13:355–361 359
123
causeway cannot grow in the high salinity of the north—is
also supported by these experimental data. N. spumigena
was found only in inocula from the less saline waters south of
the Antelope Island causeway, and apparently cannot sur-
vive the high saline waters north of the railway causeway.
Experimental support for these two general hypotheses
helps shed light on our original question: are cyanobacte-
rial distributions in the Great Salt Lake influenced by
abiotic factors, biotic factors, or both? From these experi-
ments, it appears that both abiotic (salinity) and biotic
(interspecies competition) factors seem to affect distribu-
tions of cyanobacterial species. N. spumigena distributions
seem to be primarily influenced by salinity, since it can
only grow in fresher waters. By contrast, A. halophytica
distributions seem to be primarily influenced by competi-
tion from N. spumigena and Oscillatoria sp. There are, of
course, other geochemical processes which we did not
measure but which may affect distributions. We are also
interested in the relationship between the green alga
Dunaliella and cyanobacteria. Our initial analysis suggests
a commensalism with Dunaliella benefitting from the
presence of nitrogen fixing Apanothece: in our microscopic
analysis we often observed Dunaliella cells clustered
around mass colonies of Apanothece. It would be inter-
esting if nitrogen fixed by Apanothece in hypersaline
environments contributed to exceptional salt tolerance of
Dunaliella (Zamir et al. 2004).
These experimental results are consistent with Gause’s
principle, which predicts that no two species can
indefinitely occupy the same niche (Gause 1969; Hardin
1960), since there is a clear niche partitioning between
A. halophytica and N. spumigena in
the Great Salt Lake.
These two species cannot occupy the same hypersaline
habitat north of the railway causeway, since N. spumigena
cannot tolerate hypersaline conditions, and A. halophytica
is suppressed in the presence of N. spumigena in the less
saline southern waters.
However, this leaves unanswered the question of why
A. halophytica is not totally excluded from the south, since it
occurs in all samples of inocula, regardless of salinity.
Perhaps A. halophytica is periodically excluded from
southern waters by N. spumigena blooms, but during
intervals between blooms, the extremely small A. halophytica
persists, albeit at lower levels. Thus, Gause’s Principle
should perhaps include a clarification: two species cannot
indefinitely occupy the same niche, except when that niche
is temporally partitioned, as occurs with episodic blooms of
Nodularia. This structuring of the cyanobacterial regimes by
salinity (Williams 1998) is consistent with the intermediate
salinity hypothesis of David Herbst (1999): ‘‘Abundance of
salt-tolerant organisms is limited by physiological stress at
high salinities, and by ecological factors, such as predation
and competition, in more diverse communities at low
salinities’’. Since N. spumigena distributions cannot survive
the high salinity stress of waters from the north arm of the
Great Salt Lake, and Aphanothece halophytica is competi-
tively excluded by N. spumigena at lower salinities, the
intermediate salinity hypothesis may apply.
The precise set of conditions that trigger episodic
N. spumigena blooms is unknown. Our data suggest that
these episodic blooms play a major role in excluding
A. halophytica from vast areas of the Great Salt Lake.
Being able to predict the occurrence of Nodularia blooms
would not only be of theoretical importance; it might also
lead to a better understanding of cyanobacterial blooms and
cyanotoxin impacts on wildlife and human health (Cox
et al. 2005; Metcalf et al. 2008).
Acknowledgments We thank J. Metcalf for assistance in cyano-
bacterial identification, J. Gardner for assistance in scanning electron
microscopy, and B. Schaalje for assistance with biostatistics. We are
grateful to the Wood Family Foundation for the Mus Views DIC/
Fluorescent Microscopy Facility at the Institute for Ethnomedicine,
and A. Fransiscana and R. Smithson for inspiration in our studies of
the Great Salt Lake.
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Abstract
Introduction
Materials and methods
Experimental cultures
Cyanobacterial identification
Data analysis
Results
Experimental cultures
Cyanobacterial identification
Data analysis
Discussion
Acknowledgments
References
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/ColorACSImageDict <<
/QFactor 0.76
/HSamples [2 1 1 2] /VSamples [2 1 1 2]
>>
/ColorImageDict <<
/QFactor 0.76
/HSamples [2 1 1 2] /VSamples [2 1 1 2]
>>
/JPEG2000ColorACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000ColorImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasGrayImages false
/DownsampleGrayImages true
/GrayImageDownsampleType /Bicubic
/GrayImageResolution 150
/GrayImageDepth -1
/GrayImageDownsampleThreshold 1.50000
/EncodeGrayImages true
/GrayImageFilter /DCTEncode
/AutoFilterGrayImages true
/GrayImageAutoFilterStrategy /JPEG
/GrayACSImageDict <<
/QFactor 0.76
/HSamples [2 1 1 2] /VSamples [2 1 1 2]
>>
/GrayImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/JPEG2000GrayACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000GrayImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasMonoImages false
/DownsampleMonoImages true
/MonoImageDownsampleType /Bicubic
/MonoImageResolution 600
/MonoImageDepth -1
/MonoImageDownsampleThreshold 1.50000
/EncodeMonoImages true
/MonoImageFilter /CCITTFaxEncode
/MonoImageDict <<
/K -1
>>
/AllowPSXObjects false
/PDFX1aCheck false
/PDFX3Check false
/PDFXCompliantPDFOnly false
/PDFXNoTrimBoxError true
/PDFXTrimBoxToMediaBoxOffset [
0.00000
0.00000
0.00000
0.00000
]
/PDFXSetBleedBoxToMediaBox true
/PDFXBleedBoxToTrimBoxOffset [
0.00000
0.00000
0.00000
0.00000
]
/PDFXOutputIntentProfile (None)
/PDFXOutputCondition ()
/PDFXRegistryName (http://www.color.org?)
/PDFXTrapped /False
/Description <<
/ENU
/DEU
>>
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [5952.756 8418.897]
>> setpagedevice