GENERAL
I ARTICLE
Life at High Temperatures
Ramesh Maheshwari
Amazing microorganisms thrive at high temperatures incompatible with the familiar forms of life. A few species
from hot springs or from the vicinity of hydrothermal vents
at the floor of the oceans are the basis of a multimillion
dollar industry. An entirely new form ofanimal life has been
discovered around the volcanic eruptions from the sea
floor. This animal life thrives in the permanently dark
environment only because it harbors symbiotic bacteria
that synthesize food molecules from inorganic chemicals in
the emissions. Thermophilic microorganisms are involved
in composting and humification in terrestrial habitats.
Ramesh Maheshwari is at
the Department of
Biochemistry, lISe,
Bangalore. His research
interests have included
thermophilic organisms.
Introduction
For most terrestrial organisms, the optimum temperature of
growth is around 2S-30°C. Among the few exceptions in the
higher forms of life is the Pompeii worm which survives at SO°C
or more in the vicinity of geothermal vents in the deep sea and
the plant Tidestromia oblongifolia (Amaranthaceae) found in Death
Valley in California, where the hottest temperature on earth ever
recorded during 43 consecutive days in 1917 was >48 °C
(Guinness Book of WorId Records, 1999). Temperature stress is
accompanied by water stress and since all organisms need liquid
water, life at high temperatures is found only in the terrestrial
hot springs or at the bottom of ocean in the vicinity of volcanic
eruptions. This life is comprised of unicellular bacteria or organisms resembling bacteria.
Since 1960s, new discoveries keep pushing the upper temperature limit of life higher and higher (Figure 1). Because of the
lability of crucial biomolecules, for example adenosine triphosphate, ATP - an energy-rich molecule which is the principal
donor of energy in biological systems, and of nicotinamide
adenine dinucleotide phosphate, NAD(P)H - a molecule which
Keywords
Temperature, life, thermophiles, extremophiles.
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GENERAL I ARTICLE
-150- +-- Predicted upper limit for life
-140-130-120- ...-Upper temp limit for life, 2000
-110-
+--upper limit for life, 1990
-100- .
+-- Upper limit for life, 1970
90+-- Upper limit for life, 1960
+--Highest temp. Death Valley, California,JuIy 10, 1913;;
~AI Azizia, Libya, September 13,1922
........... Protoplasmic streaming ceases after 5 min heating in most e
Highest temp., Bangalore, 22 May 1931--+
0-
~
Annual average, Dalol, Denakil Depression, Ethiopia
~
Average annual temp. in tropics
Upper limit for aquatic vertebrates
{30-}
+-Topt for most animals, plants and microorganisms
-20
Average temp. of earth--+
10- +-- Tmin for E. coli and most plants
Figure 1. Thermometer of
life. The upper temperature
limit of life has been raised
higher and higher. Modified
from R Maheshwari, 2005,
Fungi: Experimental Methods in Biology. CRC Press,
Boca Raton, USA.
functions as a coenzyme in several biochemical reactions in the
cell, the upper temperature limit of life is predicted to be around
150°C, prompting researchers to search for the champion
organism.
Organisms that have an optimum temperature (Topt)of 45-50 °C
are called thermophiles. The T opt is the temperature at which
growth rate is fastest. The so-called 'hyperthermophiles' have a
T opt of 90°C or even higher. Thermophiles and hyperthermophiles have not only an elevated T max' hut also an elevated
minimum temperature of growth (Tmin)' However, the temperature range in which these organisms grow is similar to that of the
mesophiles which grow at 'normal' temperatures. This means
that the overall temperature range at which any organism can
grow is rather narrow, approximately 30°C. The T opt is always
closer to T max than it is to T min' The most heat-enduring are the
single-celled organisms which superficially resemble bacteria,
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GENERAL I ARTICLE
but are now classified in the domain of life called Archaea
(previously Archaebacteria). The differences between Archaea
and Bacteria are in the chemistry of their membrane lipids,
structure of cell walls, structure of ribosomes and growth sensitivity to antibiotics.
Lethal Effect of Heat
Why is prolonged exposure to temperatures >45 °C generally
lethal? The integrity of the cell and of the cellular compartments
known as organelles depends on the structure of membranes
which is a lipid-based sheet. Macromolecular structures depend
on the three-dimensional structure of the molecules of which
they are composed of. For example, a protein in the cell which
performs the task of catalysis of metabolic reactions or the
transport of molecules has folded polypeptide chains called the
alpha-helix. The alpha-helix is held in the helical configuration
by hydrogen bonds between the CO groups and the NH groups
four amino acids apart. DNA, the carrier of genetic information,
has the form of a double helix that is held together by hydrogen
bonds between the base pairs. Nucleic acid and protein interact
to form nucleosomes which are joined into a flexible chain.
Hydrogen bond is a weak bond, broken by heat. The threedimensional structure of macromolecules is therefore altered by
high temperatures resulting in the loss of functional properties.
We may ask why a macromolecular structure is not based on
strong covalent bonds. The answer is that cellular function is
dependent on the flexibility of a macromolecule. For example,
an enzyme (protein) molecule must be flexible for folding into a
shape into which its substrate can fit in precise orientation.
Denaturation of proteins and melting of the membranes are two
major reasons why 45°C is a sort of temperature limit for the
majority of living beings.
Life in Boiling Water
It was a great surprise to discover that a variety of single-celled
organisms, with diameter close to or less than a micron, actually
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GENERAL
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Figure 2. A boiling pool in
Yelllowstone National Park,
USA. From [1J, with permission.
require temperatures above 40-45 °C for growth. During 1965 to
1975, the American microbiologist Thomas Brock and his associates [1] discovered microbial species thriving in hot springs in
the Yellowstone National Park, USA (Figure 2). To determine if
the bacteria were actually growing in hot springs or had been
disseminated from elsewhere, the investigators immersed microscope slides in the boiling pool; exposing one side of the slide
to germicidal ultraviolet radiation at regular intervals. The
rationale being that any organism that had become attached to
the slide would be killed and not be able to reproduce. The
colony growth on the non-irradiated, but not on the control
(irradiated) side (Figure 3), proved that microbial growth was not
only occurring at temperatures close to 90°C, it was occurring at
rapid rates. The generation time in different hot springs varies
from 2 to 6 hours. In Yellowstone, water boils at 92.5°C. This
work demonstrated that life can exist at close to the boiling point
of water.
An Important Generalization in Biology
In
India,
hot
springs
(·agnikunds·) are found in
Kashmir, Himachal Pradesh,
Uttaranchal, GUjarat, w.est Bengal and in other places, but their
microbial diversity has not been
studied .
Brock investigated several types of thermal habitats. One of
these was the self-heating piles of coal-waste in the vicinity of
coal mines. Although the bulk of coal is removed, the refuse
always contains coal fragments and other organic material. Brock
discovered that this is the habitat of Thermoplasma acidophilum.
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So far, the coal-refuse pile is the only habitat where this organism has been found. Thermoplasma has no cell wall, yet can grow
at temperatures of 61°C, and at a pH close to 1 (the acidity of
O.1M hydrochloric acid). Its discovery refuted the belief that
cell-wall provides the chief protection against heat. These discoveries led to an important generalization in biology: No matter how harsh the environment, if there is liquid water, there is
life. Or to put it differently, there is no life without liquid water.
Figure 3. (left) Bacterial
growth on microscope slide
immersed in a boiling pool.
(right) No growth occurred
on the side of the slide
which was exposed to germicidal UV light at intervals.
From {1J, with permission.
Amazing Creatures in the Vicinity of Deep-Sea Hydrothermal VentslBlack Smokers
Although man had conquered the highest mountain peak, the
outer space and the moon, the greatest depths of the oceans had
remained impenetrable because of the enormous hydrostatic
pressure due to the column of water above. Only USA, France
and Japan have built submersibles capable of descending to the
ocean floor. It carries a pilot and two scientists, and a single dive
takes nearly 4 hours. The submersible is fitted with strobe lights,
portholes, and robotic arms for collecting samples. Finally, in
1977, deep sea diving to depths of some 1.6 miles (or 2.5 km) at
locations along the thick line shown in Figure 4 could be achieved.
The line represents mid-oceanic ridges where tectonic plates
that form earth's crust separate, creating fissures on the sea floor.
Sea water penetrates into the fissures and interacts with the hot,
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GENERAL I ARTICLE
Figure 4. The line represents mid-oceanic ridges
where gigantic plates that
form earth's crust move
apart, creating cracks in the
ocean floor into which sea
water seeps in and is heated
by the molten rock or
magma.
volcanic crust. The thermally-expanded, mineral-enriched water with temperature of 350°C or more rises up and exits from
chimney-like structures on the ocean floor. However, because of
the enormous hydrostatic pressure, its boiling point is raised.
The, super-heated water containing sulfides of iron and copper,
H 2S, ammonia and minerals, mixes with the cold (2-4°C) ocean
water. An amazing discovery was made in the dark deep-sea. At
places, where the water temperature is between 18 to 40°, a
variety of animals live; strikingly a tubeworm up to 3 meter long
(Figure 5), resembling a giant-sized lipstick. These animals are
not thermophilic in the sense used for the terrestrial forms.
However, hot and cold are relative terms; because the vast
volume of ocean water is ice-cold, these animals could be regarded thermophilic or thermotolerant.
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Figure 5 (left). Tube worms, Riftia pachyptila, resembling giant-sized lipstick growing near "black
smoker", a chimney-shaped vent on the sea floor spewing black 'smoke' rich in iron and sulphide
into the ocean. The super-heated hot water with dissolved inorganic nutrients gushes out and mixes
with ice-cold sea water. Strange communities of animals in symbiotic association with bacteria live
at temperature 18-40 °C. Drawing by S Mahadevan ([email protected]) based on Colleen Cavanaugh (http://ccommtechlab.msu.edu/sitesldlc-melzoolmicrobeslritiasym.html).
Figure 6(right). Certain types of bacteria living inside tube worms obtain energy by the oxidation
of reduced inorganic compounds, such as HzS, NH4, NOz' or ferrous iron. The energy (ATP) and
reducing power (NADH) are used for reactions to generate carbohydrate (CHzO) and other organic
compounds required for growth. Only chemoautotrophic bacteria are capable of doing this. From
[2}.
Since sunlight can not penetrate into the deep sea, synthesis of
organic compounds by using light energy (photosynthesis) can
not occur. This immediately raised the question, what is the
food of these strange animals? The tube worm, Riftia pachyptila,
has neither a mouth nor a stomach. This invertebrate harbors
billions of bacteria within its body cells, which utilize the
hydrogen sulfide in the geothermal emission (Figure 6). The
bacteria oxidize hydrogen sulfide and the energy derived from
the oxidation is used for the synthesis of ATP molecules and
reducing power as N AD(P)H. These molecules are required for
fixation of carbon dioxide into organic compounds by a set of
chemical reactions which are the same as in the Calvin-Benson
cycle. These reactions were first discovered in the photosyn-
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thetic green alga, Chiarella, by Calvin and associates. Melvin
Calvin was awarded the 1961 Nobel Prize in Chemistry. But here
in the dark, deep-seas life was thriving not because of photosynthesis, but chemosynthesis. In chemosynthesis, energy is obtained by the oxidation of reduced inorganic compounds, such
as NH4, N0 2, H 2 S,.or ferrous iron, and carbon by assimilation of
CO 2 • Chemosynthesis was discovered more than a century ago
by the Russian microbiologist Sergei Winogradsky in the bacterium Beggiatoa, but was not given much attention at the time.
Figure 7. Photosynthesis
versus chemosynthesis in
the ocean.
The organic compounds synthesized by chemosynthetic bacteria are used for their own multiplication and also supplied to the
host animal which provides the bacteria with raw materials,
mainly hydrogen sulphide, oxygen, and carbon dioxide, in highly
enriched concentrations. This discovery showed an entire ecosystem based on bacteria (Figure 7). The food chain starts from
the sulfur-oxidizing bacteria as the primary producers and leads
to crabs, mollusks, and tube worms as consumers. Being thermo-
Solar Energy
Ocean surface
!
~~~ hv ~~
CO2 + H20 - CH 20 + O2
Photosynthesis
Cell material
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philic is an advantage as the hot soup has high concentrations of
energy-rich sulfides in it. Therefore, most likely, the thermophiles evolved around the hydrothermal vents. Hydrogen sulphide is very poisonous. The tube worms, mussels and clams in
the deep ocean are able to avoid poisoning because the sulphur
bacteria in their body cells oxidize hydrogen sulphide to nontoxic forms. Interdependency (symbiosis) allows organisms to
live in extreme environment where each cannot live separately.
Applications of Heat-Loving Bugs
Until Brock provided a visual clue of life in hot springs, no
'sensible' microbiologist would intentionally incubate cultures
above 37°C. Thomas Brock [1] incubated a small sample of hot
spring water in a liquid medium containing salts, and fortified
with yeast extract and tryptone as carbon and energy source at
70-75 °C (temperature-enrichmentculture). A yellow, rod -shaped
bacterium had grown in the medium. This bacterium, which he
named Thermus aquaticus, has a temperature range of 50-80°C
and is widespread. It has been found also in geyser in the
bathroom. At that time no one thought that 20 years after the
discovery, this bacterium would be exploited by an American
biochemist Kary B Mullis (awarded the 1993 Nobel Prize in
chemistry) as a source of a heat-stable Taq DNA polymerase, a
most useful enzyme in biology and medical research. (In this
enzyme nomenclature, T is abbreviation for Thermus, and aq for
aquaticus). This enzyme withstands the cycles of heating and
cooling required for separating DNA strands, binding primer
DNA to complimentary sequence and for building new strands
over and over again. By this technique, called the polymerase
chain reaction or PCR, a minute (less than one nanogram)
amount of DNA can be exponentially amplified million-fold.
Kary Mullis [2] writes that he got the PCR idea during a nightlong drive in the California Mountains while his girl-friend was
sleeping in the car. Among the many applications, Taq polymerase is used in forensic medicine for establishing identities.
DNA extracted from dried blood or from sperms in vaginal
swabs can be amplified using PCR. DNA is a stable molecule; it
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has even been extracted from 18 million year old fossilized
leaves. Perhaps the controversy whether the bones and ashes
kept in a temple in Tokyo are those of Netaji Subhas Chandra
Bose can be settled by matching the DNA extracted from the
sample with that from Netaji's daughter. Steven Spielberg's
fiction movie Jurassic Park was based on revival of dead dinosaurs by DNA amplification using Taq polymerase.
Today, T. aquaticus DNA polymerase is a multimillion dollar
industry. Its patent right was bitterly fought for. Some years ago,
the prestigious British journal Nature had an advertisement by a
'biotech' company which boasted of going to the "ends of the
earth looking for novel microorganisms which may contain
useful thermostable enzymes". The 'ends of the earth' are the
bottom of the deep-oceans, and the 'useful thermostable enzymes' are the DNA polymerases having a low error rate in
copying DNA from the template DNA. Until 1999, Pyrococcus
furiosus isolated from the vent had held the record of the most
heat-loving organism. Its 'vent polymerase' (as the DNA polymerases obtained from microorganisms growing around hydrothermal vents are referred to) remains fully active following 1 hr
incubation at 95°C.
Researching in volcanic areas is a test of one's endurance. A
cartoon in Brock's book [1] shows one stressed-out Yellowstone
visitor saying to another: "Actually, with me it's not so much the
heat, it's the awful sulphur smell". Brock dedicated his book to
all those who suffered in the Yellowstone "hell". Several hot
spring waters have pH around 1.8 to 2.2. In the hot acidic pools
(pH around 1.5) Brock observed 5-10 J,Lm lobed cells adhering to
sulphur particles, and aptly named this organism Sulfolobus
acidocaldarius ('Sulfo' is latin for SUlphur, 'lobus' for lobe, 'acido'
for acid, and 'caldarius' for hot). This is an Archaean which
derives energy by oxidizing elemental sulphur to sulphate, and
can also oxidize ferrous iron. Sulfolobus has potential application
in microbial mining or microbial leaching [3], i.e., extraction of
iron from iron-containing minerals such as chalcopyrite
(CuFeS z)·
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GENERAL I ARTICLE
Thermophilic Ancestry of Life
Suggested Reading
What does the discovery oflife in hot environments tell us about
the evolution oflife? Every cell possesses genes encoding ribosomal RNA. Based on the degree of divergence of gene sequences,
a computer can arrange the sequences of ribosomal DNA into a
diagram which has the form of a tree. This tree has three
branches: the Prokaryotes (or simply Bacteria), the Archaea and
the Eukarya. The thermophilic Archaeans are at the deepest
branch of this 'evolutionary tree', reaching the hypothetical
common ancestor (progenote). Thermophiles have been found
in the fluids from borewells drilled some 4000 m deep into the
crust of earth. In the tree of life, Eukarya are connected .to
hyperthermophile root with a transition from a hyperthermophile
to thermophile to mesophile. Researchers therefore think that
our ultimate ancestor was a thermophile.
[1] T D Brock, Thermophilic
Microorganisms and Life
at High Temperatures.
Springer. Verlag, New
York, 1978.
[2] K B Mullis, The unusual
origin of PCR, Scientific
American, Vol. 262, p. 36,
1990.
[3] http://bioteach.ubc.ca/
Bioengineering/
microbialmining/
[4] H Huber, M J Hohn, R
Rachel, T Fuchs, V C
Wimmer and K 0 Stetter,
A new phylum of Archaea
represented by nanosized
hyperthermophilic sym·
bionts, Nature, Vo1.417,
pp.63·67,2002
[5] D G Cooney and R
Emerson, Thermophilic
Fungi. An account of their
biology, activities and classification, W H Freeman
and Co., 1964.
Maheshwari,
G
[6] R
Bharadwa; and M K Bhat,
Thermophilic fungi: their
physiology and enzymes,
MicrobioLMolecular BioL
Rev., VoL64, pp. 461.488,
2000.
The Smallest Organism is a Thermophile
The current record of the most heat-loving organism is by
Nanoarchaeum equitans, a nano-sized Archaean (400 nm in diameter), discovered near a hydrothermal vent off the coast of
Iceland [4]. Its T max is 121 °C, the temperature of pressurized
steam in a food cooker or in an autoclave that is used for
sterilizing. Its genome is the smallest. Not surprisingly, N.
equitans lacks genes fot synthesis of amino acids, nucleotides,
cofactors, and lipids.. ~lives·~parasitic relationship attached to
another Archaean,I~,at-'1();'98 °C, deriving metabolites
from its host. The new diScovery of Nanoarchaeum suggests that
other organisms remain to be discovered. A number of researchers in USA, Europe and Japan, which have built submersibles,
are regularly 'diving' in the deep sea to discover new thermophilic microorganisms. To accommodate the flurry of research
activity, a new journal ~mwphiles has been founded.
Microbial Thermogenesis
Most of us can only have visions of doing research around a
black smoker or a hot spring. However, some type of research on
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GENERAL
Figure 8. A mass comprising of a mixture of herbivore dung (source of nitrogen and nutrients) and hay
(source of cellulose and
hemicelluloses). The mass
of material heats up to
>50°C for several days,
favouring the growth of
thermophilic microflora
which bring about an accelerated decomposition of
material and formation of
organic manure. A similar
unwitting exploitation of
thermophilic fungi is in the
preparation of compost for
the cultivation of the edible
mushroom. A thermistor
probe connected to a battery-operated chart recorder is used to monitor
changes in temperature of
the compost.
I
ARTICLE
life at high temperatures is possible even in a backyard. It is a
common observation that when agricultural wastes or municipal refuse containing organic debris or a mixture of herbivore
dung and straw, is piled into a heap for composting, the temperature of the mass rises and its decomposition is accelerated,
yielding excellent manure (Figure 8). The heat in the heap
(Figure 9) also kills unwanted pests and pathogens. The heap of
plant material, rich in nutrients and moisture, favours the development of thermophilic microorganisms, reducing the dissipation of heat produced from the exothermic metabolic reactions
of saprophytic microbes present therein. The interior environment of a decomposing heap is therefore warm. Remarkably, the
Mallee fowl or 'the incubator bird' which is indigenous to
Australia and islands of southwestern Pacific learnt this, and
uses the heat for incubation of its eggs! It builds large mounds
(nests) by gathering forest litter and soil similar to the manmade heap shown in Figure 8. The bird feels the heat by inserting
its head and regulates the temperature in the mound (the incubator) by removing or refilling decomposing material as required (http://www.abc.net.au/science/scribblygum/October2000/
default.htm ).
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Indeed, the generation of heat from the metabolism of aerobic
microorganisms can be quite significant. It has been known
since the ancient times that agricultural products, such as hay or
food grains, stored as a heap, catch fire [5]. Early in the last
century, investigations on self-heating stacks of hay had led to
the discovery of species of thermophilic fungi (Figure 10) which
provided a link to the puzzle of spontaneous combustion. When
the moist plant material which is rich in nutrients is packed
rather loosely to allow diffusion of air, it favours the development of thermophilic microflora within. The temperature regularly rises to 60 °C, coinciding with the T max of thermophilic
fungi [6]. During one of our field visits we observed a huge pile
of smoldering b~gasse (Figure 11) near a sugar factory in
Karnataka. Although the factory-owner was unaware of a possible role of microbial thermogenesis as a cause of fire, he was
convinced that no foul play was involved; as such types of bums
had occurred before. Rather, he emphasized that fire was due to
the mass of bagasse becoming damp due to
unseasonal shower. Knowledge of life at high
temperatures provides an explanation for spontaneous combustion of stored agricultural products. Moisture is essential for the build-up of
microbial protoplasm, whereas the large size of
the mass of material is important for insulation
against dissipation of heat produced from the
exothermic metabolic reactions of the microorganisms present therein. Most likely the initial
Figure 9 (left). Thermophiles are involved in the
decomposition of a municipal garbage dump.
Figure 10 (right). Colonies
of a thermophilic fungus
(mold) growing in pure culture on nutrient agar medium at 50°C.
Figure 11. Smoldering pile
of bagasse near a sugar
factory in Karnataka.
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GENERAL
Q. Where did you get the cul-
ture of thermophilic bacterium
from?
A. From my boss.
Q. And where did he get it from?
A. From his boss in Americo.
Address for Correspondence
Ramesh Moheshwori
Department of Biochemistry
Indian Institute of Science
Bangalore 560 012. Indio
Email:
[email protected]
I ARTICLE
rise in temperature, coinciding with the T max of thermophilic
microflora, catalyzes uncharacterized chemical reactions, resulting in further heating and the mass of material catching fire.
Summary
The discoveries oudined above have demonstrated that the
study of thermophilic organisms has not only provided us with
insights on the evolution of life on this planet, but also led to a
variety of significant practical applications. More discoveries are
expected to emerge as interest on these fascinating organisms is
on the rise.
Information and Announcements
Foundation Course in Physics and Chemistry of the Earth
sponsored by Indian Academy of Sciences, Bangalore
in collaboration with University of Allahabad, Allahabad
November 7 - 27, 2005
The course is aimed at sharpening the conceptual foundations of young scientists in the Science of the
Earth, by engendering capabilities for understanding and analysing planetary systems and processes
generally and that relating to the earth in particular, in a quantitative manner. It is accordingly designed
to focus on the most basic aspects of physical and chemical principles and their numerical application in
calculating the thermodynamic conditions of critical earth processes such as lithospheric stretching
leading to development of sedimentary basins, physics and chemistry of melts, mineral phases and their
transformations through the earth's crust and mantle, amongst other topics.
Selected participants will be provided with local hospitality and actual train fare for travel by three-tier
AC or bus fare by the shortest route. Those wishing to be considered for selection to participate in the
course, should send their application to the Course Coordinator, Professor Alok K Gupta, Director,
National Centre of Experimental Mineralogy and Petrology, 14 Chatham Lines, University of Allahabad, Allahabad 211 002, Tel.0532-1150840, Fax:0532-2644951;Email: [email protected].
Applications must include the followillg ·information : (i) Name, date of birth, full postal and email
address. telephone number(s) (ii) Qualifications and nature of current activity supported by appropriate
documents (iii) a clear statement about background knowledge of the pre-requisites, and (iv) a statement
in 200 words to indicate the scientific motivation for participating in this course.
Last date for receipt of applications: September 15, 2005.
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