Environmental Microbiology (2003) 5(11), 1094–1102 doi:10.1046/j.1462-2920.2003.00509.x
© 2003 Society for Applied Microbiology and Blackwell Publishing Ltd
Blackwell Science, LtdOxford, UKEMIEnvironmental Microbiology 1462-2920Society for Applied Microbiology and Blackwell Publishing Ltd, 20035 1110941102Original Article97 k years old Halobacterium salinarum M. R. Mormile
et al.
Received 7 March, 2003; revised 24 June, 2003; accepted 24 June,
2003. *For correspondence. E-mail mmormile@umr.edu; Tel.
(+1) 573 341 6346; Fax (+1) 573 341 4821.
Isolation of Halobacterium salinarum retrieved directly
from halite brine inclusions
Melanie R. Mormile,
1
* Michelle A. Biesen,
1
M. Carmen Gutierrez,
2
Antonio Ventosa,
2
Justin B. Pavlovich,
3
Tullis C. Onstott
3
and
James K. Fredrickson
4
1
Department of Biological Sciences, University of
Missouri-Rolla, Rolla, MO 65401,USA.
2
University of Seville, 41012 Seville, Spain.
3
Princeton University, Princeton, NJ 08544, USA.
4
Pacific Northwest National Laboratory, Richland, WA
99352, USA.
Summary
Halite crystals were selected from a 186 m subsurface
core taken from the Badwater salt pan, Death Valley,
California to ascertain if halophilic Archaea and their
associated 16S rDNA can survive over several tens of
thousands of years. Using a combined microscope
microdrill/micropipette system, fluids from brine
inclusions were aseptically extracted from primary,
hopper texture, halite crystals from 8 and 85 metres
below the surface (mbls). U-Th disequilibrium dating
indicates that these halite layers were deposited at
9600 and 97 000 years before present (ybp) respec-
tively. Extracted inclusions were used for polymerase
chain reaction (PCR) analysis with haloarchaea-spe-
cific 16S rDNA primers or placed into haloarchaea
culture medium. Enrichment cultures were obtained
from 97 kyr halite crystal inclusion fluid and haloar-
chaea-containing prepared crystals (positive con-
trols), whereas inclusions from crystals of 9.6 kyr
halite and the haloarchaea-free halite crystals (nega-
tive controls) resulted in no growth. Phylogenetic
analysis (16S rDNA) of the 97 kyr isolate, designated
BBH 001, revealed a homology of 100% with Halobac-
terium salinarum. DNA–DNA hybridization experi-
ments confirmed that BBH 001 was closely related to
H. salinarum (81–75% hybridization) and its ascription
to this haloarchaea species. The described method
of retrieving particle-containing brine from fluid
inclusions offers a robust approach for assessing
the antiquity of microorganisms associated with
evaporites.
Introduction
A variety of microorganisms have been isolated from salt
deposits by enrichment of aseptically crushed halite crys-
tals (Dombrowski, 1960; Norton et al., 1993). Rigorous
surface sterilization and controls (Norton et al., 1993) pre-
cluded the possibility of laboratory contamination and
demonstrated that the isolates were endolithic. Bulk sam-
pling methods cannot determine, however, whether cul-
tured microorganisms were derived from primary or
secondary fluid inclusions or from intercrystalline grain
boundaries or intracrystalline fractures. Incontrovertible
claims of antiquity can only be made for halophiles iso-
lated from single phase, primary fluid inclusions in radio-
metrically dated salt crystals. Consequently, assertions of
viable ancient halophiles (Dombrowski, 1960; Norton
et al., 1993) have been difficult or impossible to verify.
Even with the rigorous surface sterilization procedures
and use of sterile drill bits to breach brine inclusions
(Vreeland et al., 2000), there have been a number of
concerns expressed over the integrity of the crystal stud-
ied (Hazen and Roedder, 2001) and the lack of genetic
differences between modern and ancient microorganisms
(Graur and Pupko, 2001; Nickle et al., 2002). As a means
of achieving greater specificity in the isolation of endolithic
haloarchaea from brine inclusions, this report presents the
development, evaluation, and application of a method
for extracting microorganisms from single phase, pri-
mary fluid inclusions, while under direct microscopic
surveillance.
The impetus for the development of this in situ extrac-
tion approach is twofold. First, it aims to reduce the
opportunity for biological contamination by bacteria intro-
duced into the crystal structure via microscopic fissures.
For bulk crushed salt samples, crystals must be
immersed in sterilization and wash solutions several
times for surface sterilization before enrichment. This
induces dissolution of the salt and potentially kills tar-
geted endolithic microorganisms in addition to external
microorganisms. Drilling into the interior of salt crystals
with a sterile drill bit, and extracting bacteria directly from
fluid inclusions, reduces the possibility of contamination
by exogenous bacteria. Second, the in situ extraction
approach offers enhanced specificity over the bulk sam-
ple crushing technique. In situ extraction enables the
isolation of microscopically detected cells or cell-like par-
ticles from primary fluid inclusions as opposed to har-