The Biology of HALOPHILIC BACTERIA

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The Biology of HALOPHILIC BACTERIA
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Library of Congress Cataloging-in-Publication Data
The Biology of halophilic bacteria I edited by Russell H. Vreeland,

Lawrence Hochstein.
p. em.
Includes bibliographical references and index.
ISBN Q.-8493-8841-4
I . Halophilic microorganisms. 2. Halobacterium.
3. Archaebacteria. I. Vreeland, Russell H. II. Hochstein,
Lawrence.
QR97 .S3B56 1992
589.9'5--dc20 92-18249
CIP
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INTRODUCTION
This book was written for anyone interested in learning about the biology
of halophilic bacteria. Each chapter was composed by scientists who have
spent the better part of their working lives studying halophilic bacteria. It is
not, however, a series of review articles. Each author, was asked to step
away from their special disciplinary jargon and review style in favor of a
textbook type informational overview. The intent of this book, and in fact
the entire series to follow, is to provide ecological information to molecular
biologists and vice versa.
The idea for this volume, and series, began in 1986 during a meeting in
Jerusalem, Israel. Two events at this meeting led to this concept. First, most
of the ecologists/taxonomists did not attend the "molecular" talks because
"we couldn't see the relevance". Second, during a bus tour of the Dead Sea
two molecular biologists were overheard commenting that the Dead Sea was
the only place on Earth where "nothing lives, not even bacteria". These two
scientists, incidently, had each given a paper on amino acid sequences of
purified proteins extracted from Ha/oferax (Halobacterium) marismortui which
literally means "the salt loving bacterium from the Dead Sea".
The book isn't just for scientists studying halophiles however; it is for
anyone that simply wants to learn something about the field. In addition to
providing an overview each chapter has a discussion of the relevant meth
odology. Again though, don't look here for pages of recipes or cookbook
instructions. Such things have only been included where absolutely necessary.
Instead, the methods sections provide information about why tests and assays
are performed one way and not another. The methods sections also provide
useful hints about modification of procedures, buffers, or media when work
ing with halophiles.
In the interests of space, chapters dealing with topics such as halophilic
methanogenesis, alkaline soda lakes, fungi, algae, and psychrophilic saline
environments have been reserved for future series volumes that deal specif
ically with those topics. Other topics such as algal mat communities or
intense rhodopsin biophysics have been omitted because these topics have
been extensively covered elsewhere.
The authors that contributed to this book all accepted a very difficult
assignment and did it very well, considering they had no actual format from
which to work. We want to publically thank them for their diligence and
their patience. We believe that they have developed a useful uniquely readable
volume. We hope that you, the reader, agree.
Russell H. Vreeland
Lawrence I. Hochstein
THE EDITORS
Russell H. Vreeland is an Assistant Professor at West Chester Univer
sity in West Chester, Pennsylvania.
Dr. Vreeland received a B.S . degree in Environmental Science from
Rutgers College of Agriculture and Environmental Science (now Cook Col
lege) in 1973 and an M.S. degree in Microbial Ecology from Rutgers Uni
versity in 1976. He received a Ph.D. in Microbial Physiology from the
University of Nebraska - Lincoln in 1980. Following 2 years of Post doctoral
research at the University of Western Ontario in London Ontario, Canada he
was an Assistant Professor of Biology at the University of New Orleans. He
moved to West Chester in 1989.
Dr. Vreeland has studied halophilic bacteria since 1976 when he isolated
and identified the genus Halomonas. He is a 3-term member of the Inter
national Judicial Commission Subcommittee on the Taxonomy of Halobac
teriaceae. He is the chairman of the International Working Party to Establish
Minimum Standards for the Taxonomy of Halophilic Bacteria. He is a mem
ber of the American Society for Microbiology, the American Association for
the Advancement of Science and the United Methodist Church.
Dr. Vreeland has received research grants and contracts from Sandia
National Laboratories, the U.S. Army Corps of Engineers, the Louisiana
Fisheries Initiative, and private industry. He has published 20 research articles
and reviews. Dr. Vreeland has presented numerous seminars and invited
lectures in the United States and abroad. His current research interests involve
the isolation and characterization of halophilic bacteria from hypersaline lakes
and underground salt formations .
Lawrence I. Hochstein, received his Doctorate in Bacteriology in 1959

from the University of Southern California. He was a Research Associate in
the Department of Medical Microbiology, School of Medicine, University of
Southern California, Research Associate in the Department of Genetics,
School of Medicine, Stanford University, Instructor in the Department of
Microbiology, School of Medicine, University of Illinois, and is currently
Research Scientist, Space Sciences Division, Ames Research Center, Moffett
Field, California. His major research interests include the physiology of the
halophilic bacteria, denitrification, particularly in the exteremey halophilic
bacteria, and the ATPases in the Archaea.
TABLE OF CONTENTS
Chapter 1 Introduction to Saline Environments . . . . . . . . . . . . . . . . . . . . . . . . . 1

Francisco Rodriguez-Valera
Chapter 2 Ecology of Extremely Halophilic Microorganisms .......... 25

Aharon Oren
Chapter 3 Ecology of Moderately Halophilic Bacteria ......... ........ 55

A. Ramos-Cormenzana
Chapter 4 Growth and Nutrition of Halophilic Bacteria... ..... .. . ..... 87

D. J. Kushner
Chapter 5 Taxonomy of Halophilic Bacteria .......................... 105

Russell H. Vreeland
Chapter 6 Lipids of Extreme Halophiles .............................. 135

Masahiro Kamekura
Chapter 7 Lipids of Halophilic and Halotolerant

Microorganisms ............... . ........... ...... ...... ..... 163
N. J. Russell
Chapter 8 Osmotic Adaptation in Halophilic and

Halotolerant Microorganisms .............................. 211
Johannes F. Imhoff
Chapter 9 The Molecular Biology of Halophilic

Archaebacteria ............................................. 255
Patrick P. Dennis
Chapter 10 Bioenergetics and Transport in Extreme Halophiles ........ 289

Janos K. Lanyi
Index .................................................................... 311

Chapter
1
Introduction to
Saline Environments
Francisco Rodriguez-Valera
Departamento de Genetica Molecular y Microbiologfa
Universidad de Alicante
Alicante, Spain
1. INTRODUCTION TO SALINE ENVIRONMENTS
This book is concerned with microorganisms adapted to life in an extreme

environment containing inorganic salts so highly soluble in water that high
concentrations can be reached. This is an important environmental factor on
Earth; we have only to consider that there are 4.29 X 10 16 tons of dissolved
salts in the waters of the ocean (Ronov, 1968) to realize the importance.
Moreover, over the history of the planet, these salts have been precipitated
and deposited, forming evaporite rocks amounting to 2% of the sedimentary
rock of the continents (Kirkland and Evans, 1981 ). So many of the environ
ments on our planet challenge their inhabitants with the presence of inorganic
salts of differing chemical nature, but sodium chloride (NaCl; common table
salt) is normally the most abundant. Thus, most organisms are able to deal
with moderate concentrations of salts dissolved in the water surrounding
them. Obviously, all marine organisms can endure, and very often require,
the salinity of the ocean, which is remarkably constant around the world and
is close to 3.5%* of total salts . What happens, however, when the salts
0.8493·8841-4193/$0.00 +$.SO
Cl 1993 by CRC Pl'oss. Inc. I
2 The Biology of Halophilic Bacteria
exceed these limits? A domain is entered in which salinity plays a dominant

role in determining the biota present. The environment becomes extreme and
inhibitory for the majority of organisms, so that a highly specialized micro
flora develops having a physiology favored by the presence of high concen
trations of salts, mostly NaCI. These organisms are the halophiles. This
chapter considers the saline environments, their special characteristics and
how they interact with halophilic microorganisms. As stated, hypersaline
environments are extreme environments, so the chapter begins by analyzing
the concept of extreme environments and extremophiles.
2. EXTREME ENVIRONMENTS
A simple definition of extreme environments is that they are unfavorable,
or even lethal, to most living organisms. "Most" is emphasized because
organisms exist that are specifically geared to a life in extreme environmental
conditions. These organisms, recently named extremophiles, not only tolerate
but prefer or even require extreme conditions for their development. Brock
(1979) devised a more precise definition, based on taxonomic criteria: "en
vironments in which species diversity is low, and some taxonomic groups
are missing''. Low taxonomic diversity is a feature common to all extreme
environments. Usually a gradient can be found, diversity decreasing as
conditions become more extreme (Rodriguez-Valera, 1988). These extremely
simple ecosystems are ideal to study, and the development of models of
theoretical ecology, nutrient cycling, and trophic chains can be very simple.
What conditions make an environment extreme? Almost any factor af
fecting biological processes can be responsible for creating extreme environ
ments. Life is widely distributed on the Earth's surface (hydrosphere and
atmosphere); however, certain conditions are required in order to perform
the physiological processes that are essential conditions for the maintainance
of a stable ecosystem. One oft-recognized condition is the existence of liquid
water. This does not imply that the temperature range for life is between 0°
and 100°C. Cells can accumulate solutes and lower the freezing point several
degrees. At the other extreme, pressure can keep water liquid at very high
temperatures, as happens in deep ocean hydrothermal vents. Brock (1979)
hypothesized that life is possible over the whole range of temperatures where
water remains in liquid form, but recent evidence indicates that the range is
probably much narrower (White, 1984; McMeekin and Franzmann, 1988).
Another basic condition for life is the existence of a source of biologically
utilizable energy and nutrients for the building of biomass. The Earth's
environments that meet these conditions are diverse, however, most have the
following characteristics: intermediate temperatures, neutral pH, pressure of
only a few atmospheres, abundant oxygen, and the salinity of fresh- or
seawater. These are what could be called "normal conditions" in the sense
3
that they are very common in the biosphere. It is, therefore, quite under
standable that the evolution of most organisms has caused them to live in
these conditions and to be affected negatively by important deviations from
these values.
It is obvious that life processes are strongly affected by the environment;
the macromolecules essential to life, as well as the chemical reactions oc
curring in biological processes, are in general rather sensitive to physico
chemical parameters. It is also true that living cells have ways of creating
an intracellular environment that can be very different from the milieu
outside. However, the capacity of cells to control physical parameters such
as temperature, pressure, or radiation is very limited in all but highly evolved
multicellular organisms. Moreover, certain essential structures and enzymes
(e.g., cell envelopes or transport systems) must be exposed. As a result,
every organism has its own optimum, maximum, and minimum environ
mental conditions for life. The response in microorganisms is usually mea
sured by growth, which is simple to quantify. The growth range is useful
because it shows the set of environments in which the organism can play an
active role. Growth response and survival are also extremely diverse in
microorganisms, and the growth range of organisms isolated from different
environments often do not overlap. For example, the temperature range for
growth of bacteria from thermal and cold environments can be as much as
100°C apart, The shape of the response curve can also vary from wide to
narrow, with a steep slope and narrow maximum or with a relatively ho
mogeneous, rather flat response.
This diversity reflects the plasticity of life. Enzymes, membranes, and
nucleic acids can be adapted to fulfill their functions under very different
conditions. When the nature growth range of one organism moves into an
environment apart from the conditions preferred by most others (an environ
mental extreme), the organism becomes an extremophile. Most extremophiles
become confined to extreme environments because the adaptation process is
usually accompanied by a loss of the capacity to grow in normal conditions.
What kind of evolutionary trend can drive an organism to become an
extremophile? There may be different answers to this question. First, we
have to consider that the predominant conditions on Earth have changed
throughout its history. What is extreme now could have been normal in the
Archaean, and organisms that evolved in those times could still thrive,
utilizing pockets of primitive conditions remaining on the planet. It has been
speculated that one extremophilic group (the archaebacteria) (Woese, 1977)
belongs to this type. All the archaebacteria inhabit environments which can
be considered extreme in some aspects - thermal, hypersaline, or anaerobic
- and they are also the most extreme of extremophiles, the most thermo
philic, halophilic, and oxygen-sensitive organisms. Also, their extremophilic
physiology is of a type that it is difficult to explain as having evolved from
a "normophilic" ancestor.
Another type of extremophile is the kind that has evolved from an

organism living in a normal environment, or perhaps a more moderate one.
It seems logical that such extremophiles would develop the simplest physi
ological mechanisms of resistance, or those requiring the minimum amount
of change of genetic information (Yancey et al., 1982), producing less
specialization.
As stated by Brock (1979), "only extreme environments able to serve
as sites for long-term evolution, will allow the development of complex
mechanisms of adaptation to their conditions". I would add that the long
term would also allow relatively more different species (phylogenetic lines)
to adapt. Obviously, there is also an effect inherent to each specific envi
ronmental factor, and the degree to which it disturbs biological processes,
on the amount of time required for evolution to produce an organism adapted
to the specific set of conditions.
In summary, we can imagine an infinite number of extreme environ
ments, since almost any factor will influence biological processes. Only
those that are present in nature in some abundance, however, will be of
biological interest and will have produced specialized extremophiles.
3. TYPES OF SALINE ENVIRONMENTS
There are two major types of biologically important environments in

which the salt factor will interact with the microbial populations, soil and
water. Soils containing >0.2% soluble salts are considered saline (Kaurichev,
1980). They are very common around the world, particularly (although by
no means exclusively) in arid regions. Many plants are adapted to develop
under these conditions (globally termed halophytes). The range of salinities
that can be found in soils is very wide and a complex microflora develops
over most of this range (Quesada et al., 1982); however, the existing infor
mation about halophilic soil microflora is very fragmentary.
Saline waters are much better known. Besides the ocean proper, there
are many waters with a salinity similar to that of seawater. Both coastal
systems, such as salt marshes, and some saline lakes far from any coast can
be included. Lakes are considered saline if they have >0.3% salinity (De
Dekker, 1983). Very few halophilic organisms are found in this type of
water. Hypersaline waters are those containing salt concentrations notably
higher than those of seawater (Edgerton and Brimblecome, 1981 ). Since both
the concentrations of salts and their nature can vary, such hypersaline systems
can be classified according to different criteria.
There are two major factors to consider; the first is the origin and the
nature of the salts. Many hypersaline bodies of water arise from the evapo
ration of seawater, and such aquatic environments are called thalassohaline
(from the Greek thalasso, the sea). In a strict sense thalassohaline waters are
5
those related by origin to the sea, the most typical example being a coastal
lagoon fed by seawater. Nevertheless, the term is normally used to refer to
any hypersaline water body in which salts are present in roughly the same
proportions as those found in seawater (Brock, 1979; Williams, 1981 ). This
loose delimitation includes many environments with salts originating from
mineral deposits, but where chemical compositions are similar to those of
brines produced by the concentration of seawater. In this sense the term can
include all chloride-sulfate waters (see below). For instance, the Great Salt
Lake, located nearly 1000 km from the Pacific Coast is thalassohaline. From
an ecological point of view, the concept of thalassic waters is consistent
because the organisms and ecological interactions seem to be similar in all
such environments.
Other hypersaline waters derive from the dissolution of salts of conti
nental origin. Many of these lack some of the components found in seawater
salts, or contain different predominant ions, varying notably from the pro
portions in seawater. These are called athalassohaline waters. The compo
sition of athalassohaline waters depends on many factors (discussed later),
which are accordingly very diverse, although some biological or chemical
patterns are more common than others. In the most common types special
populations of organisms with specific adaptations can be found . In others,
due to their special characteristics a normal thalassic microflora is present,
though diversity is often restricted.
The second factor to consider when classifying hypersaline waters is the
salt concentration. The lower limit to which the term hypersaline can be
applied is not clear, and different authors apply different arbitrary limits
ranging from 3 to 12% salinity (Post et al., 1983; Por, 1980). One argument
which has often been used is that in hypersaline waters the high concentration
of salts is the prime factor in the drastic reduction of species diversity (Por,
1980). In other works, a limit is located at around lO to 12% salinity, below
which other environmental factors predominate in determining the popula
tions of organisms; above it the salinity factor becomes the major determinant
and the biota found is similar regardless of geographical location. This lO
to 12% range is sometimes called mesosaline (Kirkland and Evans, 1981).
For thalassohaline, there is also an upper limit of concentration that can be
established at around 35%. Here, NaCI starts precipitating copiously and the
proportions of salts start shifting rapidly to a magnesium preponderance.
However, the concentrations at which NaCI precipitates also marks the upper
limit of resistance of all biological forms (Javor, 1983).
4. ORIGIN, DISTRIBUTION, AND

ABUNDANCE OF TYPES
As previously mentioned, the distribution and abundance of a specific

type of extreme environment is very important for the degree of specialization
of the biota. If one type of environment is common in the biosphere, has a

wide geographical distribution, and possesses a certain constancy in its
characteristics throughout geological periods, one can presume that a long
and complex evolutionary process could have taken place. In the case of
hypersalinity it is clear that the above considerations apply to thalassohaline
waters. The alkaline-carbonate lakes also possibly conform to these condi
tions. In answering the question of distribution and abundance of hypersaline
waters, it is essential to consider their origin-under what conditions, and
how the hypersaline water bodies arose.
The place of origin of the salts found in most hypersaline environments
is seawater. The II cardinal ions of oceanic water are Na+, K+, Mg 2 +,
Ca2 +, Sr+, CI-, SO/-, HC03 -, Br-, p- and H3B03 • Chloride (55.04%)
and sulfate (7.68%) are the predominant anions, and sodium (30.61%) and
magnesium (3.69%) the predominant cations (Pipkin et al., 1977). Salinity
is extremely constant in the present-day ocean, with an average salinity of
3.5% and a normal variation of only ±0.2% (Macintyre, 1970). Whether
the concentration and types of salts dissolved in the ocean have changed
much throughout the Earth's history is a matter of controversy. It is possible
that Na + did not predominate in the early Archaean and the quantities of
Mg 2 +, Ca2 +, and K+ could once have been similar to Na+ (Ronov, 1968).
In any event, the salinity of the ocean has been nearly constant for the past
200 million years (Macintyre, 1970).
When seawater becomes separated from the open ocean evaporation
causes the concentration of the salts, while rain- and freshwater flowing into
the basin dilute them. If the net balance is in favor of concentration the water
will eventually become hypersaline. The first salt to precipitate in important
quantities is calcium sulfate (gypsum), although some carbonates will pre
cipitate earlier in small amounts. Gypsum will start to precipitate when
seawater reaches about one third of the original volume (approximately 10%
salinity). Halite (NaCl) will only start to precipitate when seawater is reduced
to one tenth the original volume (34% salinity). Finally, Mg 2 + and K+ salts
precipitate around l/20th the original volume (Blatt et al., 1980). However,
slight changes in local environmental conditions will give rise to considerable
shifts in the relative amounts of various salts crystallized. In fact, some 54
salts, double salts, and hydrated salts can be obtained by evaporating seawater
(Macintyre, 1970).
The salts deposited during this evaporation process are the origin of
evaporite rocks. Such rocks can be considered imprints of ancient hypersaline
water bodies and show the stability and extent of some of them. It must be
borne in mind that evaporating a column of normal seawater 1000 m deep
to dryness will produce only 0. 75 m of solid gypsum and approximately
13.7 m of halite (Blatt et al., 1980). The deposition ofthick strata of evaporite
minerals usually implies that new salts are continuously provided for precip
itation in the form of new seawater entering through a narrow inlet or by
7
seepage from the sea. The large amounts of gypsum and halite present in
ancient formations show that depositional environments often existed for very
long periods of time. New salts were continuously supplied by new saline
water entering the system. A similar phenomenon occurs today in the so
called "sabkhas", an Ethiopian word that refers to a marine lagoon in which
new seawater penetrates by seepage from the sea or by inflow through a
narrow inlet, and is evaporated to the point of precipitation of the different
salts.
Although evaporite depositional environments are required for the pre
cipitation of the large amounts of salts often found in these rocks, other
hypersaline environments derived from seawater evaporation exist in consid
erable abundance along the shorelines of arid and semiarid zones. These
natural salines are very common on flat coasts with low pluviosity. Human
activity has transformed many of these salt pans into the solar salterns that
are widely distributed around the world.
Some solar salterns have a single, large pond in which a volume of
seawater is evaporated; however, most consist of many small ponds con
nected by canals and ditches. The first ponds are filled with seawater; when
the salt concentration increases slightly, it is permitted to flow or is pumped
to the next pond(s). The salts concentrate due to evaporation as the water
circulates throughout the system. Finally, NaCl is precipitated in special
ponds (crystallizers). This system offers an excellent model of aquatic en
vironments with different salinities. They have been studied by a number of
authors and provided very interesting information about the effect of salinity
on the microbial populations that develop (Rodriguez-Valera et al., 1981,
1985; Javor, 1983a, b).
Another possible origin of hypersaline waters is the dissolution of salts
from rocks by continental or meteoric waters. The salts are usually concen
trated further by evaporation once they accumulate in an impermeable basin.
This kind of environment is as diverse as it is abundant and widespread. The
chemical composition of some closed basin lakes is determined by the pres
ence of mineral deposits in the lake basin. Obviously the nature of the rocks
from which the salts were dissolved also influences the composition of the
brine. Some chloride can also come from ocean-derived rainwater, and thus
the chloride content is influenced by the distance between the basin and the
ocean (Brock, 1979). However, the precipitation processes occurring while
evaporation takes place are essential in determining the final composition,
and these are extremely complex and difficult to predict (Harvie et al.,
1980). Each saline lake is therefore a special case; however, they can be
classified into two major types depending on the predominant anions, the
chloride-sulfate lakes, and carbonate-chloride lakes. It seems that although
many elements are involved in deciding which type will develop, a key factor
is the availability of divalent Ca2 + and Mg2+ cations. When Ca2+ and Mg2+
Figure I . Distribution of some well-known hypersaline waters of the world, according to the
anionic composition of their waters. The two main types of saline waters are clearly recognizable.
(From Por, F. D., Mar. Ecol. 1:121-131, 1980. With permission.)
are scarce a carbonate-chloride lake that is high in pH will develop that will
force the precipitation of small amounts of these divalent cations.
When the divalent cations are more abundant, as in seawater, carbonate
ions precipitate and a chloride-sulfate lake results. These lakes have neutral
pH and sometimes, as with the Great Salt Lake, they are thalassohaline.
When a chloride-sulfate brine is confined in a reducing environment,
such as when large amounts of organic matter are available with little aera
tion, sulfate is reduced to sulfide and in the presence of high Mg2 + and Ca2 +
will produce a neutral to slightly acidic lake such as the Dead Sea (Ordoiiez,
1984). A common expression of the chemical composition of closed-basin
lakes is by means of a triangular coordinate plot, in which the relative anionic
composition of Cl - , SO/-, and C032 - are given at the apex of the triangle.
Figure 1 shows the distribution of some hypersaline lakes according to the
anionic composition of their waters. The two main types of saline waters
(chloride-sulfate and carbonate-chloride) are clearly recognizable.
5. BIOLOGICAL CONSIDERATIONS:
HALOTOLERANT AND HALOPHILIC ORGANISMS
Any organism living in hypersaline environments must cope with a
number of environmental stresses related to the high ionic concentration of
the medium. The presence of high amounts of electrolytes in the medium
produces several effects. First, there is a decrease in water activity, i.e., the
9
amount of water available for biological processes. In this respect salinity

resembles desiccation and this explains, for example, the similarity between
plants from desert and saline environments. On the other hand, salts are
efficient osmolytes and create an osmotic imbalance to cells containing
normal levels of cytoplasmic solutes. Lipid membranes allow rapid diffusion
of water molecules in and out of cells while presenting an effective barrier
to most other biological molecules (Csonka, 1989). This situation implies
the loss of water and an irreversible retraction of the cytoplasm (plasmolysis)
(Rose, 1976). This is the speediest lethal effect exerted by high salinity and
is probably responsible for the rapid killing effect of concentrated salt solu
tions on microorganisms. A more subtle but no less important effect of
concentrated salt brines is dependent on the chemical nature of the ions,
since they exert destabilizing effects on proteins and lipids.
The effects of salts on proteins have long been known. In 1888, Hof
meister classified cations and anions as "salting-out" or "salting-in" by
their effect on the solubility of proteins and other colloids (Yancey et al. ,
1982). Salting-out ions decrease the solubility of polar molecules in water.
These salts strengthen hydrophobic bonds in proteins. Salting-in salts have
the opposite effect (Lanyi, 1974). Electrostatic forces among amino-acyl
residues are also altered in the presence of salts. All these effects have major
consequences on the tertiary and quaternary structure of proteins which are
essential for their biological activity.
Lipid bilayers are also affected dramatically in electrolyte solutions
(Quinn, 1986). The hydrophobic and hydrophilic interactions responsible for
the stability of lipid bilayers are altered. Also, electrolytes may cause de
hydration of membranes by competing with membrane polar groups for free
water. Some solutes may partition into the hydrophobic domain of the mem
brane and destroy long-range interaction of the hydrocarbon region. All of
the complex biophysical consequences of the presence of salts in an envi
ronment require profound adaptation at the molecular level to allow required
biochemical processes to function in concentrated salt solutions.
In addition, an organism adapting to life in hypersaline habitats, in which
salt concentrations change over wide ranges, must be able to function over
those ranges. In other words, the capacity for osmoregulation is an important
characteristic of inhabitants of hypersaline environments. As with other en
vironmental variables the ability to grow at different salt concentrations is
varied in microorganisms (Figure 2). Most freshwater and marine microor
ganisms are rather stenohaline, with a sharp growth optimum around their
natural salt content. However, some organisms whose usual habitat is not
hypersaline can withstand relatively high salt concentrations, albeit with a
reduction of the growth rates . This occurs in halotolerant microorganisms.
Some halotolerant microorganisms can grow at high salt concentrations, and
often can be found in hypersaline habitats. They tolerate hypersaline con
centrations but they are still negatively affected by salts; their adaptation is
incomplete.
PR-6 Fs Ds Ah H
•
0
IX
.t:
j
I!)
::
0 •te SDl
0 M NoCI I 3 4
Figure 2. Growth response to salinity for five organisms. PR-6, AgmeMllum qUlldraplicatum
(Cyanobacteria). Ab, Aphanot~c~ ha/ophitica (Cyanobacteria). H, Halobact~rium sp. Fs, Fabr~a
salina (protozoa). Ds, Dunaliella salina (algae). (From Post, F. J. et al., Hydrobiologia 105:95-
113, 1983. With permission.)
A different case is that of halophiles, which show more or less wide

ranges of growth at different salinities, but experience optimum growth (as
measured by the growth rate) at salinities significantly higher than seawater.
In other words, halophiles are specialized for living in hypersaline environ
ments and their specialization implies a dependence on salt for optimal
growth, and, very often, even for survival. The concepts of halophilic and
halotolerant organisms have been subjected to different interpretations by
different authors (Larsen, 1986; Kushner, 1978; Vreeland, 1987). Many
consider the need of salt for survival essential for an organism to be consid
ered halophilic (Larsen, 1986). However, there are examples of bacteria that
can grow without any salt added to their culture medium, but which still
require concentrations twice that of seawater to grow optimally. (A. Ventosa,
personal communication). There are also some technical problems related to
the description of the salt-growth response .
One of the technical problems is the mixture of salts considered. All
known halophilic organisms are adapted to life in environments in which
NaCI is the major component, but other salts are also present and they can
markedly affect the response of the organism to NaCI. The mixture of salts
present in seawater and which remain in solution as it concentrates can be
considered a single ecological factor. For organisms from thalassohaline
environments the ideal mixture for measuring the salt response may be
different concentrations of this mixture mimicking different natural environ
ments. Obviously, this is not applicable to organisms from athalassohaline
waters. Another aspect to keep in mind when studying the salt response is
the influence of other environmental factors, particularly temperature (Post,
1977) and nutrients present (Forsyth and Kushner, 1970). The salt concen
tration influences the optimum growth temperature and vice versa. Thus, it
is necessary to provide the temperature at which a salt response curve has
been developed, and this should always be close to the optimum temperature.
In addition, a rich medium should be used to avoid inhibition of growth due
11
to the higher nutritional requirements sometimes associated with high salinity

in the medium (Forsyth and Kushner, 1970).
Halophilic microorganisms are relatively diverse, being found among
the eubacteria, archaebacteria, and eukaryotes. They are also very cosmo
politan: the microbial populations found in hypersaline waters of similar
concentrations have the same composition regardless of geographical location
or even climatological conditions (Por, 1980). Only in Australian saline lakes
do there seem to be some cases of endemism, apparently due to the long
isolation of this continent (De Dekker, 1983). Even in Australia, the more
concentrated brines have the microbial flora characteristic of other hypersa
line waters.
What kind of modifications must an organism undergo to become hal
ophilic? All halophilic organisms have developed specific adaptations to live
in high salt concentrations. All have an efficient mechanism for osmoregu
lation, adapting their intracellular concentration of osmolytes to the external
salt concentration so that the net osmotic imbalance is small (Brown, 1976).
For this purpose most halophilic microorganisms use neutral organic com
pounds called compatible solutes (Brown, 1976) that cause very little alter
ation to, or even protect, the enzymes and other components of the cytoplasm
(Mackay et al., 1984). An important exception to this is halobacteria, and
also some anaerobic halophilic eubacteria (Oren et al., 1984), which accu
mulate inorganic compatible solutes. Second, all organisms have the means
to exclude toxic ions such as sodium or chloride from the cytoplasm. Third,
all halophilic organisms have adapted their external structures, cell wall,
membranes, and extracellular and membrane enzymes to work partially or
totally exposed to the high salt medium. In the halobacteria this adaptation
applies also to cytoplasmic components that are exposed to extremely high
concentrations of potassium, sodium, and chloride.
Some of the above-mentioned physiological mechanisms for coping with
high salt concentrations are not exclusive to halophiles, i.e., osmoregulation
is probably present in all microorganisms and therefore the only change in
halophiles is the scale of the process. The compatible solutes found in
halophilic eubacteria, for example, are also found in their nonhalophilic
counterparts (Le Rudulier and Bouillard, 1983). The exclusion of certain
ions from the cytoplasm is also not exclusive to halophiles, and is probably
just a matter of more efficient transport of these ions against stronger gra
dients. Of the points mentioned above, the third must represent a longer and
more complex evolutionary process involving changes of the sequences of
many proteins, metabolic pathways, and of synthesis of structural molecules.
This third aspect of the adaptation also explains the salt dependence of
most halophiles. Extracellular materials, membrane proteins, and lipids are
irreversibly adapted to high salt. There are indications that the adaptation of
proteins to work in high salt medium involves profound changes. As men
tioned above, the electrostatic and hydrophobic bonds responsible for the
final three-dimensional structure of proteins are strongly affected by a high

ionic strength (Lanyi, 1974). The presence of salting-out salts encourages
the formation of hydrophobic interactions, an effect similar to increased
temperature. The enzymes of the halobacteria are stabilized by some of the
salts, according to the Hofmeister series (SO/- > CI- >> N0 3 - and Na+
> K+ > Cs +) (Werber et al., 1986).
Another central problem for enzymes exposed to high salt is how these
proteins can retain a sufficient amount of hydration. Halophilic enzymes of
both halobacteria (Reistad, 1970) and some halophilic eubacteria (Imhoff,
1986) have a high proportion of acidic aspartate and glutamate but a low
content of hydrophobic and basic amino acids. This partially explains the
high levels of hydration and salt binding found in most halophilic enzymes
(Werber et al., 1986). To obtain such enzymes from nonhalophilic precursors
would likely involve a long and tortuous evolutionary process.
6. CHARACTERISTICS OF HYPERSALINE
ENVIRONMENTS
What are hypersaline environments like with respect to other environ

mental factors relevant to the biota? This is a very complex question due to
the diversity of this type of environment. The following paragraphs review
the tendencies in the different hypersaline environments that have been
studied.
6.1. Temperature
Although arid tropical climates with high average temperatures are the
regions in which hypersaline habitats are most abundant, saline lakes of
different origins are also found in temperate and even polar latitudes. Under
conditions of high light intensity and high daytime temperatures, waters with
high salinity heat up very rapidly because of a combination of physical and
biological factors. Concentrated salt solutions have low specific heat and
evaporation rates (Copeland, 1967), and the pigmentation of microorganisms
present in the water increases heat absorption by acting as an infrared trap
(Davis, 1978). As a consequence, very high temperatures can be reached,
even with mild air temperatures. In Mediterranean salterns, for instance,
maximum temperatures of 45°C have been registered in air temperatures of
30°C (Rodriguez-Valera et al., 1985). Conversely, hypersaline waters cool
faster at night or during the winter, and remain liquid at very low tempera
tures. In the Great Salt Lake temperatures range from - 30°C in winter to
+48°C in summer (Post, 1981). Some hypersaline lakes were recently de
scribed in Antarctica that maintain low temperatures year round . Organic
Lake, which has a salinity that varies between 15 and 21 %, has temperatures
13
ranging from -14° to + l5°C (Franzmann et al., 1987). A similar range is

found in Deep Lake with a salinity of 28% (Barker, 1981). To summarize,
most hypersaline water bodies are subject to high temperatures, with large
periodic fluctuations. Some permanently cold hypersaline lakes are also
known.
The vast majority of halophilic microorganisms are mesophilic or slightly
thermophilic. However, some halophilic bacteria with a moderate psychro
philic response have been isolated from the Antarctica hypersaline lakes
(McMeekin and Franzmann, 1988). Generally speaking, halophiles have a
preference for high temperatures. In the Great Salt Lake, for instance, aerobic
heterotrophs have optima ranging between 37° and 50°C. The most extensive
bacterial growth occurs in summer near the shore where the highest temper
atures are reached (Post, 1977). Halophilic archaebacteria have optima be
tween 40° and 55°C (Juez, 1988). Their enzymes are also markedly ther
mophilic. Halobacterium menadione reductase, for example, has a maximum
thermal stability close to 85°C, among the highest found in any protein
(Borowitzka, 1981 ). Nitrate reductase from H. salinarium has a maximum
activity at 25° to 30°C, but only in the presence of high salt concentration.
It has been suggested that high salt concentrations such as those found in the
cytoplasm of halophilic archaebacteria confer thermal stability to halophile
proteins because they promote the establishment of hydrophobic interactions
within the proteins (Lanyi, 1974). In this sense, halophilic bacteria show a
peculiar connection between temperature and salt growth response, increasing
their salt requirements at high temperatures. This happens both in halophilic
archaebacteria and eubacteria. One possible explanation is that both temper
ature and salinity produce similar modifications of the properties of water,
such as the size of molecular aggregates. Therefore, high salinity and low
temperatures could act synergistically to inhibit metabolic processes (Lanyi,
1974). In halophilic eubacteria there are also examples of slight thermophil
ism. For example, most halophilic photosynthetic anaerobes of the genus
Ectothiorhodospira have temperature optima between 45° and 50°C (Ray
mond and Sistrom, 1969; Imhoff and Triiper, 1977). The high temperature
tolerance in these organisms could be explained by the presence of the
compatible solutes, mostly betaine (Galinski and Triiper, 1982), that act as
enzyme protectants.
The halophilic alga of the genus Dunaliella, although mesophilic, has
considerable resistance to both high and low temperatures which is attributed
to the large intracellular concentration of glycerol that serves to reduce the
effects of heat and cold stress on enzymes (Borowitkza, 1981 ). Microbial
populations have also been found in the cold lakes in Antarctica. In Deep
Lake, there is a Dunaliella sp., and a species of halobacteria (Halobacterium
lacusprofundi) that can grow, although extremely slowly, at 4°C. The growth
response to the temperature of this species is about 5°C below that of species
from warm climates (McMeekin and Franzmann, 1988). A halophilic eubac-
teria, Halomonas subglaciescola, with a growth range between -4° and

+ 32°C, has also been isolated from Organic Lake (Franzmann et al., 1987).
6.2. Pressure
Most hypersaline lakes are shallow and therefore develop no great pres
sure at the bottom. However, there are some important exceptions. The
deepest part of the Dead Sea (320 m) where considerable anaerobic life has
been detected is subjected to considerable pressure (Nissenbaum, 1975).
Density stratification tends to originate more concentrated brines at the
bottom of meromictic lakes (Burton, 1981).
The Red Sea geothermal brines are a peculiar example of deep hyper
saline waters. These brines, located at the bottom of the Red Sea, are formed
by Red Sea water which leaches through evaporitic facies, then rises and
discharges through vents into two deep basins (about 2000 m) at the bottom,
about 1000 km further north, in the central rift of the Red Sea (Craig, 1969).
These brines are composed of evaporite salts-mostly halite (NaCl), but also
gypsum and anhydride (CaS04 ), with a final salinity of 28% and a compo
sition similar to seawater, although with lower concentrations of Mg 2 + and
sulfate. The water also contains significant amounts of heavy metals and is
heated geothermal1y, reaching 56°C in one basin and 45°C in the other
(Simoneit et al., 1987). This is an example of a large body of water (the
layer of brines in the larger basin has a surface of 75 km 2 and is several
meters deep) subjected to a number of simultaneous environmental stresses:
high salinity, temperature, heavy metals, and pressure. Sulfate reduction of
bacterial origin was found in one of the basins (Discovery Deep) (Watson
and Waterbury, 1969) and the other was presumed to be sterile. Unfortu
nately, no extensive microbiological work was carried out with samples from
these environments, which are among the most extreme natural environments
on Earth.
6.3. Availability of Nutrients

The presence of organic and inorganic nutrients affects the nature of the
biota. As a general rule, hypersaline waters tend to be very productive
(Kirkland and Evans, 1981; Post, 1977). The large number of indigenous
consumers, such as some bivalves and/or flamingos, is a reflection of this
high productivity. Spectacular blooms of Dunaliella and/or halobacteria are
also indicative of the extremely high productivities present in the higher
salinity range which reflects the availability of phosphorus and nitrogen. The
net influx of seawater from which evaporite depositional environments of
the sabkha types originate acts as a continual supply of allocthonous nutrients.
The shallowness of most hypersaline water bodies facilitates a rapid recycling
of nutrients which do not accumulate on the bottom, unless there is a great
proliferation of microbial mats (see below).
15
Reports abound about the nutrient contents in different hypersaline waters.

Post (1977) concluded that in the Great Salt Lake phosphate was plentiful at
the sediment surface. Orthophosphate remained fairly constant, declining
slightly during algal blooms. Nitrogen, either as ammonia, nitrates, or ni
trites, was low or undetectable. He concluded that the Great Salt Lake was
limited in inorganic nitrogen. By contrast, in the Dead Sea the limiting
nutrient is phosphate, being undetectable in the water, while ammonia (2 to
6 mg/1) and organic nitrogen (0.3 to 3 mg/1) are abundant (Kaplan and
Friedman, 1970). In marine salterns the situation is different and at times
contradictory. Javor (l983b) describes two such marine salterns. In one
saltern, located in Baja California (Guerrero Negro), the nutrient concentra
tions and the primary productivity were very low. Javor (1983b) attributed
this to the development of extensive benthic microbial mats in the first stages
of the brine concentration. These mats are presumed to sequester the nutrients
and leave nutrient-poor overlying water. The other pattern described corre
sponded to a saltern located near San Diego, CA. Nutrient concentrations
and plankton productivity were higher while the benthic communities were
much more restricted . This latter case seems to be the more common.
In some multipond marine salterns the concentration of nitrogen and
phosphorus increased with the salt concentration, as did the biomass, reaching
the highest values in the crystallizer ponds where NaCI precipitates (Javor,
1983; Rodriguez-Valera et al., 1985; Rodriguez-Valera, 1988; C. D. Litch
field, personal communication). Alkaline saline lakes are also characterized
by high productivity. Imhoff et al. (1979) reported nitrogen and phosphate
to be in abundance in the Wadi Natrun lakes, and the productivity of these
lakes is very high. In fact, some of the highest productivity values found in
phytoplankton in aquatic environments have been reported for alkaline saline
lakes (Tailing et al., 1973).
6.4. Light
Hypersaline waters are often located in geographical regions with few
cloudy or raindays per annum. They tend to be shallow. Therefore, the biota
is exposed to strong solar radiation, and some of the most characteristic
inhabitants of hypersaline waters are red due to carotenoids that serve as
protective agents. In Dunaliella salina an interesting correlation is found
between pigmentation and the salt concentration of the medium, since higher
concentrations of salt induce the accumulation of carotenoids .
Light also plays an important role in the biology of halobacteria, where
it is used by bacteriorhodopsin and other light-driven ion pumps (Lanyi,
1984). Other retinal-proteins permit a complex phototactic behavior. The
peculiar carotenoids of the halobacteria, bacterioruberins, act primarily as
photoprotectants, but other roles have been suggested. For instance, they are
involved in an energy transfer capacity that facilitates photoreactivation, via
visible light, of DNA damaged by exposure to ultraviolet (UV) radiation

(Hescox and Carlberg, 1972). It has also been suggested that the red color
of halobacteria acts as an infrared (IR) trap increasing the heat accumulation
of the water which is favorable to these slightly thennophilic organisms
(Larsen, 1980). In some halobacteria the production of carotenoids is also
affected by the salinity of the medium, but in a direction opposite of that of
Dunaliella salina. Some species of halobacteria belonging to the genus
Haloferax grow optimally with around 20% salts in the medium, but at this
concentration only small amounts of carotenoids are synthesized. However,
at suboptimal salt concentrations, close to the lower limit of the salt-growth
range (10% salts), the cells are strongly pigmented (Kushwaha et al., 1982).
Whether these effects are caused by the salt sensitivity of the enzymes
involved in carotenoid synthesis or whether they have a specific biological
role is not known.
6.5. Oxygen
Since high salt concentrations and high temperatures restrict the solubility
of oxygen, hypersaline waters have low concentrations of oxygen (Copeland,
1967); however, aerobic life is well represented in these environments. In
shallow ponds or lakes with high algal or cyanobacterial productivity the
oxygen transfer rate to the aerobic microbial populations can be high enough
to permit a relatively rapid metabolism. Alternatively, the small storage
capacity and low diffusion rate of oxygen in brines would easily permit the
development of localized anoxic conditions. The high density of the brines
is another factor that contributes to the easy generation of anaerobiosis by
favoring stratification. Some halobacteria possess gas vacuoles which could
help to maintain them near the surface where the oxygen concentration is
highest.
7. METHODS FOR STUDY AND FIELD WORK

The study of hypersaline habitats is in most respects similar to that of
other aquatic or soil environments. A knowledge of the geological and
climatological setting is particularly important in this case because these
aspects can considerably affect salinity fluctuations and changes in the salt
composition of the water. For example, whether the rainfall is homogene
ously distributed throughout the year or concentrated in one season can affect
the biota, with a preponderance of more euryhaline types in the latter case.
This is particularly true for hypersaline soil, where the rain causes much
more drastic changes in the salinity of the microhabitat of soil particles
(Quesada et al., 1982). In order to describe a hypersaline ecosystem, as with
any other ecosystem, both physicochemical features and biological compo
sition must be established. I will consider the first type of factors, since the
others are discussed in other chapters of this book.
17
7.1. Salinity
Which salt species are present and at what concentration is obviously an
aspect of paramount importance. In thalassohaline waters derived from the
evaporation of seawater the determination of the qualitative composition of
the brine may not be necessary; however, the exclusively marine origin of
such brines must be demonstrated, since even coastal hypersaline lagoons
may have surface or groundwater inlets that change the final salt proportions.
As is done with seawater the salt concentrations of pure thalassohaline waters
and salts can be roughly inferred from the chloride concentration. Chloride
can be determined by simple silver nitrate titration. It must also be remem
bered that calcium and carbonates precipitate very early in the concentration
process. Possibly the simplest method to determine total salinity in a sample
is measuring the dry weight, e.g., drying at ll0°C to weight constancy. Salt
is very hygroscopic, so precautions must be taken to avoid hydration of the
samples during weighing. With the above method organic matter is included
in the weight. In brines with low biomass this is not normally a very important
error. If necessary, it can be eliminated by heating the salts to 400°C, after
which only inorganic material is left (including the ashes of the organisms
persent). Some methods of measuring salinity can be carried out in situ,
which can be very convenient. One such method is measuring conductivity.
However, with concentrated brines the use of normal conductimeters requires
a precise dilution of the sample. An extremely easy and reliable method for
measuring salinity is by means of a Baume hydrometer. This is a simple
device that measures water density by immersion. The ratio between density
given in Baume degrees (0 8e) and grams per cubic centimeter is d = 146.78/
146.78 - Be. An optical diffractometer that measures the diffraction index
of the water is another way of estimating salinity in the field . Some com
mercially available instruments give a direct value for the salinity of the
water.
To properly characterize the salts present in a sample and their quantities,
however, it is necessary to resort to standard chemical analysis. Flame and
atomic absorption spectrophotometry and ion analysis by liquid chromatog
raphy, or direct coupled plasma analysis, have greatly simplified many of
these analyses. Indirect coupled plasma analyzers should not be used with
concentrated brines since salt precipitates will clog the orifices, causing errors
and expensive cleaning.
7.2. Salinity of Soils

In agricultural science soils that contain >0.2% soluble salts are consid
ered saline (Kaurichev, 1980). For a proper description of the salinity of
soils an analysis of the composition of anions (C0 3 - . HC0 3 - , Cl , S04 - )
and cations (Ca2 · , Mg2+, Na +, K +) is necessary. A simple determination of
the soluble dry residue can be used to give an indication of the total salinity
in the soil. In a typical procedure, 100 g of dried soil are shaken with 500
ml of C02-free water for 3 min. The suspension should be filtered through
paper until a clear filtrate is obtained, then the dry weight determined as
described above (Kaurichev, 1980).
7.3. Other Physicochemical Determinations

Many chemical determinations are affected by high concentrations of
salts. This factor should be considered when performing any analysis of
saline samples. Since enzymes from nonhalophilic organisms function sub
optimally at high concentrations of salts, enzymatic methods should take this
effect into account.
The methods used for the analysis of seawater can be employed for most
chemical determinations. Filtration of high salinity samples can be a problem
due to the density of the concentrated brines and the presence of microscopic
NaCl crystals. Fiberglass filters work better and should be used whenever
possible. For the filtration of large volumes of highly saline water tangential
filtration systems give excellent results.
8. EVOLUTION OF THE HALOPHILIC

CHARACTER: HALOPHILES AND THE
EVOLUTION OF SALINE ENVIRONMENTS
Halophilism has arisen separately in all the major groups of microorgan
isms (Table 1). There are halophilic representatives in eukaryotes, algae,
protozoa, and fungi. There are also halophilic members in at least two of the
major branches of archaebacteria, methanogens, and halobacteria (Larsen,
1986). Most of the major phylogenetic branches of eubacteria, including
purple bacteria, spirochaetes, cyanobacteria and Gram positive, have halo
philic species (Ventosa, 1988).
Within each of the three urkingdoms there seems to be a tendency to
use the same types of organic material as compatible solutes. Betaine, for
instance, is widely distributed in eubacteria (Imhoff and Rodriguez-Valera,
1984), and glycerol in eukaryotes (Borowitzka, 1981). The archaebacteria,
on the other hand, seem to eschew organic compatible solutes, opting instead
for accumulating K+ intracellularly. There are interesting exceptions, such
as the presence of betaine in halophilic plants and methanogens, or the
accumulation of K+ by anaerobic, halophilic eubacteria.
It therefore seems possible that the basic osmoregulatory systems were
established after the separation of the three urkingdoms and before their
diversification into major phylogenetic branches. Convergent evolution based
on the particular advantages of the use of certain compatible solutes such as
19
TABLE 1.
Halophilic Microorganisms Among the Different
Phylogenetic Branches of Microbes
Eubacteria
+ Gram-positive
+ Purple bacteria-Pseudomonas
+ Spirochaetes
+ Bacteroides-Fiavobacterium-Cytophoga +
+ Cyanobacteria
Green sulfur bacteria
Green nonsulfur bacteria
Sulfate reducers, myxobacteria
Radioresistant micrococci
Planctomyces
+ Haloanaerobiaceae
Eucaryotes
+Fungi
+ Protozoa
+Algae
Archaeobacteria
+ Halobacteria
+ Methanogens
S-dependent
betaine or glycerol is a possible alternative (Yancey et al., 1982). It is also

possible that the common ancestor of the three urkingdoms used a mixture
of compatible solutes, including some now used by more advanced organ
isms.
K+, however, is the most likely candidate involved in primitive osmo
regulation. All cells have the ability to concentrate K + against a gradient.
In archaebacteria the presence of a high intracellular concentration of K +
seems to be common, even in nonhalophilic representatives (Sprott and
Jarrell, 1981). It has been suggested that the origin of life occurred near
shore tidal pools where organic matter was concentrated by evaporation
(Nisbet, 1985). The salinity of such environments would be expected to be
high. In this case, the common ancestor could have been halophilic, sug
gesting that halophilic communities existed since the initial steps of evolution.
Halophilic biota are known to aid in the development of hypersaline
habitats. At lower concentrations the microbial mats make the bottom of the
ponds impermeable, limiting the loss of concentrated water from the basin
(Davis, 1978). At higher concentrations the bright red color of many halo
philic microorganisms increases heat absorption in the brines, expediting the
evaporation processes. Lovelock (1988) has hypothesized that living organ
isms play an important role in creating evaporitic environments and hence
in regulating the salt concentration of the oceans. He specifically credits the
halophilic biota with a role in preserving new salt deposits by forming a coat
over the layers deposited. This avoids or at least retards the dissolution of
such salt deposits by rain. Certainly, it would be difficult to imagine evap
orative lagoons without living organisms. The sodium ion and the other
positive ions of K +, Mg2+, and Ca2 • all have relatively short residence
times in the ocean. They are removed by biochemical and chemical processes,
as well as by hydrothennal chemical reactions within seafloor spreading
zones, and deposited as sediments, clays, limestones, and dolomite. How
ever, chloride is quite unreactive. There is no significant biochemical trade
in chlorine (Lovelock, 1988). The salt concentration of the ocean is in a
subtle equilibrium, and evaporitic environments act as a chloride sink that
decreases the concentration of this toxic anion. Salt is continuously added to
and removed from the oceans. Some of this salt is deposited in evaporite
beds and subsequently buried beneath sediments. These deposits are tem
porary stores that the movement of the Earth and weathering continuously
expose and release back to the sea. By aiding the efficiency of these systems,
halophilic organisms could play an essential role in keeping the aquatic
environments of the Earth moderate with respect to salt concentrations (Lov
elock, 1988).
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The Biology of HALOPHILIC BACTERIA

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The Biology of halophilic bacteria I edited by Russell H. Vreeland,

Developed by Telford Press

C> 1993 by CRC Press, Inc.

International Standard Book Number Q.-8493-8841-4

Library of Congress Card Number 92-18249

Printed in the United States of America I 2 3 4 5 6 7 8 9 0

Printed on acid-free paper

Lawrence I. Hochstein, received his Doctorate in Bacteriology in 1959

Chapter 1 Introduction to Saline Environments . . . . . . . . . . . . . . . . . . . . . . . . . 1

Chapter 2 Ecology of Extremely Halophilic Microorganisms .......... 25

Chapter 3 Ecology of Moderately Halophilic Bacteria ......... ........ 55

Chapter 4 Growth and Nutrition of Halophilic Bacteria... ..... .. . ..... 87

Chapter 5 Taxonomy of Halophilic Bacteria .......................... 105

Chapter 6 Lipids of Extreme Halophiles .............................. 135

Chapter 7 Lipids of Halophilic and Halotolerant

Chapter 8 Osmotic Adaptation in Halophilic and

Chapter 9 The Molecular Biology of Halophilic

Chapter 10 Bioenergetics and Transport in Extreme Halophiles ........ 289

Index .................................................................... 311

1. INTRODUCTION TO SALINE ENVIRONMENTS

This book is concerned with microorganisms adapted to life in an extreme

exceed these limits? A domain is entered in which salinity plays a dominant

Another type of extremophile is the kind that has evolved from an

3. TYPES OF SALINE ENVIRONMENTS

There are two major types of biologically important environments in

4. ORIGIN, DISTRIBUTION, AND

As previously mentioned, the distribution and abundance of a specific

of the biota. If one type of environment is common in the biosphere, has a

amount of water available for biological processes. In this respect salinity

A different case is that of halophiles, which show more or less wide

to the higher nutritional requirements sometimes associated with high salinity

final three-dimensional structure of proteins are strongly affected by a high

What are hypersaline environments like with respect to other environ­

ranging from -14° to + l5°C (Franzmann et al., 1987). A similar range is

teria, Halomonas subglaciescola, with a growth range between -4° and

6.3. Availability of Nutrients

Reports abound about the nutrient contents in different hypersaline waters.

visible light, of DNA damaged by exposure to ultraviolet (UV) radiation

7. METHODS FOR STUDY AND FIELD WORK

7.2. Salinity of Soils

7.3. Other Physicochemical Determinations

8. EVOLUTION OF THE HALOPHILIC

betaine or glycerol is a possible alternative (Yancey et al., 1982). It is also

of Methanospirillum hungatei and Methanobacterium thermoautotrophicum. Can. J.

study. J. Bacteriol. 166:559-564.

organic solute production and excretion by the yeasts Hansenula

studies on the internal solutes of xerophilic fungi. J. Gen. Microbiol. 129:2915-2925.

Jovanovich, S. B., M. T. Record, Jr., and R. R. Burgess. 1989. In an Escherichia

moderately halophilic bacterium. Can. J. Microbiol. 19:1181 -1186.

their pathways in Escherichia coli. Eur. J. Biochem. 151:613-619.

of Methanospirillum hungatei and Methanobacterium thermoautotrophicum. Can. J.

Biophys. Acta 559:377-397.

archaebacterium Haloferax volcanii. Arch. Microbiol. 151:530-536.

in Halobacterium halobium differs from F0F1-ATPase/synthase. J. Biochem.

vesicles of Halobacterium halobium. J. Biochem. (Tokyo) 103:231-236.

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What are hypersaline environments like with respect to other environ