Mammoth Hot Springs, Yellowstone National Park, 1982 |
I wrote and submitted a memo to
Director Hat Yoder requesting permission to spend four months studying and
collecting thermophiles in Yellowstone National Park:
“Carbon
isotope ratios of naturally occurring blue-green algae have yet to be measured
that show a similar fractionation to the Precambrian carbon. Some of these
modern stromatolites consist primarily of photosynthetic, mat-forming bacteria,
Chloroflexus. Perhaps the difference between modern blue-green algal δ13C and
Precambrian δ13C is that the photosynthetic bacteria, not the blue-green algae, were
responsible for the stromatolites found in the Precambrian.”
No one had
measured the isotopic compositions of any of these organisms before. The idea
that different carbon assimilation and metabolic pathways might impart
different isotope fractionations was an exciting new lead. Yoder liked to keep a “tight ship” and to
know what his staff was doing at all times. Every year in April, Director Yoder
sent out a memo to staff members detailing the timeline for writing and
producing the Carnegie Institution’s Annual Yearbook Report. We termed this the
“rigid adherence” memo because in the memo’s first paragraph Yoder stated
“Rigid adherence to the schedule” was required. I had to promise to handle all
of my Yearbook submissions prior to departing for the west. Somewhat
reluctantly, Hat Yoder approved my request. Armed with a permit from
Yellowstone National Park (YNP), my first big field season began.
Carnegie scientists had previously
worked in Yellowstone National Park. In 1935, E.T. Allen and Arthur Day
published a megavolume on the chemistry and location of all of the thermal
springs. Allen and Day consulted with a University of California botanist W. A.
Setchell to learn about the organisms in the hot springs they were studying:
“Accepting
the facts of biological research, the writers concede that algae are a factor
favoring the precipitation of both travertine and silica from these thermal
waters. Professor W. A. Setchell concludes from his observation in the
Yellowstone Park and elsewhere, that a few species of blue-green algae are
probably instrumental in the formation of limited amounts of both travertine
and silica deposits, though they are not responsible for anywhere near all of
either. Dr. Setchell lays special emphasis on several species of algae, within
whose woolly or gelatinous layers masses of travertine, shaped by the growing
plant, accumulate. These particular organisms may indeed exercise a more potent
influence than the rest; still, from the chemical viewpoint, all
photo-synthetic plants, so far as they derive their carbon dioxide from the
water, should be regarded as a factor in the formation of carbonate.”
My field plans included sampling a
series of hot springs with very different water chemistries. I highlighted
carbonate springs vs. silica depositing springs vs. highly acidic springs.
Brock and others had shown that even in nearly boiling water at pH 2.0,
organisms could survive and even thrive (e.g. Sulfolobus). For my first
year of fieldwork, I had to choose hot springs that were within 1 to 5 km of a
road as well as relatively hidden from Park visitors. At this time, park
regulations stipulated that the public should not view researchers in action, a
policy that has changed since then. The silica depositing springs I chose were
in the Lower Geyser Basin: Fairy Creek, Queen’s Laundry, Wiegert’s Channel, and
Octopus Spring. As part of this work, I incubated glass slides in the spring to
sample freshly grown material that matched with the environmental conditions
that I was measuring at the same time (e.g., temperature, pH, CO2 concentration). Once sufficient growth was noted on the glass slides,
biomass was scraped off and dried for isotopic analysis.
There were three major photosynthetic
organisms in the neutral pH, silica depositing springs: Synochocccus lividus
(coccoid, yellow cyanobacteria); Chloroflexus sp. (a phototrophic
eubacteria); and Phormidium sp. (orange, filamentous conophyton-forming
cyanobacteria). Rarely were these organisms found in isolated forms, however,
temperature segregated the populations. S. lividus grew at the highest
temperatures (60-75°C); Chloroflexus slightly lower (48-65°C); Phormidium
at the lowest temperatures (40-48°C). The pink filaments from Octopus Spring,
were identified in the 1990s as Aquifex species (Reysenbach, 2012 ).
In the early 1980s, the carbon fixation
pathways in many of these organisms were not known. In the ensuing decades,
detailed pathways for the diversity of microbial carbon fixation were
discovered. Chloroflexus, for example, uses the 3-hydroxyproprionate
cycle, whereas Aquifex sp. uses the reductive TCA cycle. Cyanobacteria
all use Rubisco and the pentose phosphate pathway for CO2 fixation. Unwittingly, I had been measuring microbes with three
different CO2 fixation pathways with attendant carbon
isotope signatures. House et al. (2003) published a paper summarizing
literature data from cultures, along with original data from culture
experiments. Organisms using the 3-hydroxyproprionate pathway had the smallest carbon
isotope fractionations averaging around; those associated with the reverse TCA
cycle were intermediate; and organisms using the pentose phosphate pathway, had
the largest fractionation patterns.
Without knowing the biochemistry behind these different pathways, I
discovered them through measuring their carbon isotope signatures. The work
applies directly to understanding the carbon isotope patterns that we find in
the Earth’s oldest rocks.
From this project, I learned how to
carry out fieldwork in a remote, distant location where one had to be
resourceful and calculating. There were grizzly bears in the Park, bison and
elk that commonly charged tourists, and of course, boiling hot springs with
fragile crusts. The work required permitting, regulations, and reporting. The
park supervisor required that researchers do their work out of the public view,
something very difficult in crowded and popular areas. I chose sample locations
off the beaten track. Taking samples and setting up biogeochemical measurements
in the field, preserving the samples properly for subsequent isotopic analysis,
and creating experimental protocols were all skills learned during the three
summers in Yellowstone and carried through my years as a field savvy
biogeochemist.
One of the major findings of my studies
at Yellowstone was discovering the relationship between the concentration of
inorganic CO2 with the carbon isotope patterns of
organic carbon fixed by cyanobacteria and other photosynthetic microorganisms
(Estep, 1984). The relationship between the carbon isotopes and carbon dioxide
concentration was not well known at the time. I hypothesized that high
concentrations of CO2 in the Precambrian would have resulted in
more greater isotope differences because Rubsico would have an unlimited supply
of carbon for fixation. Organisms collected in acidic springs (pH 4-5) plotted
on a different line, probably because the inorganic carbon source was CO2 rather than the bicarbonate ion at this pH. Around the same time, Farquhar, O’Leary, and
Berry (1982) published their equations describing how intra- and extracellular
partial pressures of CO2 influence carbon isotope fractionation
in higher plants. Subsequently, such relationships have proven to be important
for interpreting paleo-CO2 conditions by measuring compounds from
living organisms found in marine sediments (see Freeman, 2001, for a review).
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