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JASWANT SINGH
AND
H.N. DUTTA
EDITORS
CONTENTS
Foreword
vii
Preface
ix
xi
Contributors
xiii
Acknowledgments
xvii
Chapter 1
Chapter 2
13
Land-Ice-Air-Ocean Interactions
in the Schirmacher Oasis, East Antarctica
Khwairakpam Gajananda1 , H. N. Dutta2 and Victor E Lagun3
39
89
Chapter 3
Chapter 4
Chapter 5
Chapter 6
Chapter 7
Chapter 8
107
131
149
163
vi
Contents
Sanjeeva Nayaka1 , Dalip K. Upreti1 and Ruchi Singh2
Chapter 9
Chapter 10
Chapter 11
Chapter 12
Index
173
199
221
233
249
FOREWORD
The Indian Antarctic program started in the year 1981-82 and soon it was realized that
Antarctica holds strongly coupled ice-air-ocean interactive system in the world. Dr. Ram
Manohar Lohia Avadh University had an opportunity to participate in four Indian Scientific
Expeditions to Antarctica in collaboration with the National Physical Laboratory, New Delhi.
I am happy to see the results from the various Indian Antarctic expeditions being poured
in the form of a book, which deals with a complex subject of ice-air-ocean interaction over
the icy continent. I am also happy to note that there are contributions from Italian and Russian
Antarctic stations supporting the SCAR spirit.
Antarctica holds the most efficiently coupled ice-air-ocean interaction system which has
led to many unique natural phenomena both over the Southern Ocean and over the Antarctic
continent itself. In fact, it is the efficient coupling in Antarctica that would help it to maintain
and sustain the much debated global change in various ways.
I am sure; the book will fill up the gap of scientific results particularly from the east
Antarctic regions and the understanding on the ice-air-ocean interactive processes over the
Antarctic continent and would inspire the younger generation to formulate their projects for
future expeditions to Antarctica from the world over.
As an academician, I compliment the authors (Dr. Jaswant Singh & Dr. H.N. Dutta) and
all the Authors of various chapters for integrating their knowledge for the advancement of
Antarctic science & technology. I wish authors would continue their efforts in future in
elevating Antarctic research for the benefit of humanity, which is facing multiplicity of
complex problems at the global level.
(Prof. R.C. Saraswat)
PREFACE
Antarctica is the coldest, windiest, driest and the shiniest continent on Earth, which is
surrounded by the stormiest and biologically the most productive oceans in the world. To a
first time visitor, Antarctica is seen like a fantasyland and an ivory vastness bisected by
ethereal mountains. In addition, this place contains many scientific mysteries like aurora,
polar shadows, mirage, katabatic winds, severe cyclones, ozone hole and many other
geophysical phenomena. The continent covers an area of 14x106 km which is about 10% of
the land surface on earth. Antarctica holds a pristine environment and the world's largest
stock of fresh water in the form of a frozen thick ice sheet. Antarctica pushes ice, cold fresh
water and cold air mass from its interior towards the Southern Ocean in order to maintain cool
around it and to lead many atmospheric/oceanic processes. But the ocean around Antarctica
transports back only the air laden with rich water vapor and some minor constituents present
in the air. The water vapor condenses to cause snowfall both over the Antarctic continent and
over the ocean itself. The snowfall not only maintains the equilibrium for Antarctica to retain
its physical strength and shape, it is absolutely necessary to maintain pristine environment.
Antarctic environment is the least polluted and is having the highest atmospheric visibility.
Antarctica is a magnificent display of interaction between air and the various
phases of water in a pristine environment. This interaction has led to the
formation of many unique features and many scientific mysteries, which are yet to be
explored. Antarctica is now emerging as an important key in the understanding of global
environmental concerns. Its unique features have provided scientists with special
opportunities to investigate the origin of the continents, the pollution of the globe, ozone
hole and changes in world climate. Lack of scientific data remains a major problem for
researchers in many areas of Antarctic science; it is basically due to vast size of the continent,
inhospitable conditions and the logistic support to sustain the efforts.
The contributors are eminent scientists, expedition members and technologists, who have
directly experienced the vastness of this continent to realize the Mother Natures
craftsmanship in carving the most efficiently, coupled system of ice-air-ocean in the world.
The book shall be useful to almost all the planners and researchers working in the area
of Antarctic science and technology as it encompasses chapters specifically devoted to
Antarctic science, land-ocean-ice-air interaction, influence of solar wind on atmospheric
processes, atmospheric observation at Dome C, navigation with global positioning system in
Preface
Antarctic circle, effects of UV-B radiations on terrestrial ecosystem, Antarctic lichen and
mosses and adaptations of Antarctic flora to survive under extreme environmental conditions.
Advances made in recent years have been provided in this book which will be helpful in
understanding of Antarctic climate and environmental changes. The book provides up-to-date
information on Antarctic ice-air-ocean environment. It is hoped that the book shall benefit the
polar scientific community in general and shall stimulate further advancement in the polar
science.
CONTRIBUTORS
Anand K. Dubey
Department of Environmental Sciences, Dr. R.M.L. Avadh University, Faizabad224001, U.P., India.
Prof. A. K. Gwal
Space Science Laboratory, Department of Physics, Barkatullah University, Bhopal
462026, India.
Dr. Dalip K. Upreti
Lichenology Laboratory, National Botanical Research Institute, Rana Pratap Marg,
Lucknow 226001, U.P., India.
Prof. Dipl.-Ing. Nitosh Kumar Brahma
Department of Chemical Engineering, Indian Institute of Technology, Kharagpur721302, W.Bengal, India.
Dr. H. N. Dutta
Shri Balwant Institute of Technology, Sonepat-131001, India.
Dr. I. Pietroni
ISAC-CNR via del Fosso del Cavaliere, 100, 00133 Roma, Italy.
Dr. Jaswant Singh
Department of Environmental Sciences, Dr. R.M.L. Avadh University, Faizabad-224001,
U.P. India.
Dr. Khwairakpam Gajananda
Department of Environmental Science, Faculty of Science, Addis Ababa University,P.O.
Box 1176,Addis Ababa, Ethiopia
Dr. L. V. Egorova
Arctic and Antarctic Research Institute, St. Petersburg, 199397, Russia.
xiv
Contributors
Dr. Laxmi Bishnoi,
Government Girl's P. G. College, Gurgaon-122 001, India.
Dr. N.C. Deb
Indian Statistical Institute, 203 B.T. Road, Kolkata-700 108, India.
Dr. O. A. Troshichev
Arctic and Antarctic Research Institute, St. Petersburg, 199397, Russia.
Dr. Pawan K. Sharma
Department of Chemistry, Kurukshetra University, Kurukshetra-132 119, India.
Dr. P. K. Purohit
National Institute of Technical Teachers Training and Research, Shamla Hills, Bhopal462026, India.
Dr. Rajesh Tiwari
Electrical, Electronics and Computer Engineering
Newcastle University, Newcastle Upon Tyne, NE1 7RU, UK.
Rudra P. Singh
Department of Environmental Sciences, Dr. R.M.L. Avadh University, Faizabad224001, India.
Dr. Ruchi Singh
Plant Physiology Laboratory, National Botanical Research Institute, Rana Pratap Marg,
Lucknow 226001, U.P., India.
Sanghdeep Gautam
Department of Environmental Sciences, Dr. R.M.L. Avadh University, Faizabad224001, India.
Dr. Sanjeeva Nayaka
Lichenology Laboratory, National Botanical Research Institute, Rana Pratap Marg,
Lucknow 226001, U.P., India.
Contributors
xv
ACKNOWLEDGMENTS
On behalf of all the authors, the editors are thankful to the National Centre for Antarctic
and Ocean Research, Goa (Ministry of Earth Sciences, Government of India, New Delhi) for
providing us an opportunity to participate in various Indian Scientific Expeditions to
Antarctica. These endeavors have given us a personal experience to bring before the world a
volume of scientific work for a better understanding of the ice-ocean-air interactions over
Antarctica.
The editors would also like to express their sincere thanks to the authorities of the
National Physical Laboratory, (Council of Scientific and Industrial Research, New Delhi) and
Dr. R.M.L. Avadh University, Faizabad, U.P. for wholeheartedly supporting the Indian
Antarctic program and our personal participation.
The editors would like to express thanks all the authors who have provided their scientific
contributions in the form of various chapters. The views expressed in this book are solely of
authors and are not to be attributed to their respective governments and institutions.
In this series of acknowledgements, we sincerely express gratefulness to our families for
their moral and emotional support while the work was being carried out over the icy
continent.
The editors particularly wish to thank Nova Science Publishers, Inc. and their staff
members for their collective efforts in finalizing and publishing this book. At the end, we
would definitely like to thank our readers and we are sure, there might be many shortcomings
but we will be very happy to receive suggestions and the criticism to improve ourselves.
Editors
Chapter 1
INTRODUCTION
The universe has seven continents namely Asia, Africa, North America, South America,
Antarctica, Europe and Australia. Among all continents on the basis of shape and size
Antarctica occupies the fifth place. Antarctica is the remotest, isolated and frozen continent of
the Earth, with a very inhospitable terrain and a very harsh climate. Antarctica is arguably the
most untouched region on the planet that makes it one of the world's most important places to
do scientific research. Prolonged stay and research on this continent is fraught with dangers.
Antarctica being called as the heaven for scientist and has several unsolved mysteries
concealed within it. It is a universally accepted fact that this is the largest pollution free
research lab available to the mankind. Antarctica occupies an area between South Pole and
Antarctic Circle from 65S and 90S. Antarctic convergence zone lies southward from 50
south and comprises of all islands presents beyond 50 but the actual zone is beyond 66.5
south. This island is banked by water on all sides being Atlantic Ocean, Pacific Ocean and
Indian Ocean. Being the highest, coldest, driest, windiest continent it is engulfed by ice to
almost 97.6 % and the rest 2.4% part of this continent comprises of iceless part of land.
Almost forming one- tenth part of the earth and its area is 14million sq. km. The features of
suspense and thrill possessed by this continent fascinate the world. Antarctica is also known
as the land of midnight sun, most unbearable weather, house of hail and snowstorms, land of
the penguins, a frozen desert, fascinating continent and South Pole with all its attractive
features attracts one and all.
Two hundred million years ago Antarctica was joined with Africa, Australia, India, New
Zealand and South America forming the super continent- Gondwanaland. Forces within the
Earth affecting the crust caused these continents to separate and drift apart. Thirty million
years ago the continents as we know them today, reached their approximate present positions.
Humans didn't even catch a glimpse of Antarctica until 250 years ago. And only in the last 90
years have people begun to explore this vast polar desert in earnest. After conquering the
South Pole and the discovery of Antarctica this land happens to be the centre of interest for
the scientists to quench their aspirations, and this land no longer remained lonesome. Initially
Antarctica was not a frozen land around 70 million years ago and the environment was as
similar to other warm islands. There were flora and fauna on the land. But little is known
about it today. A new scientific era has already begun with the discovery of Antarctica.
India has been sending scientific expeditions to Antarctica every year since 1981-82, and
has collected a wealth of data in various areas of science. With the establishment of Dakshin
Gangotri in 1983-84 and Maitri in 1988-89, India is now directly involved in polar research.
The fossils of the past atmospheric inhabitants lie in the layers of ice. Due to excess of ice
the plant samples are well protected waiting to be unearthed. Antarctica is a vital area of
attraction not only due to its scientific value but also due to its geopolitical and commercial
values. Its comprises of valuable minerals like copper, gold, zinc, manganese, tin, uranium
various gases and oil. But the mining of minerals and commercial usage are not allowed at
present. The water bodies have species that may possess some commercial usage. Nature has
bestowed on the Antarctica some ornamental and natural blessing, which are of prime
importance due to their geographical and scientific values. It is very surprising to enlist that
the ice of Antarctica holds about 90% of pure water of the world. Several such icebergs melt
here every year. Scientists are making an analysis of expenditure regarding transportation of
these icebergs by huge ships to fulfill the requirements of drinking water. A single small
iceberg has the potential to fulfill the water requirement of a metro town in prime countries.
An iceberg that broke form this continent in 1963 is perhaps the largest ever known iceberg
with a length of 335 kms and width being 97 kms. It floated for 12 years before losing its
existence.
In the interior regions extremely low temperatures, several months of complete darkness,
fierce winds and blowing snow combine to make life virtually impossible. Antarctica is the
harshest in the world. Wind chills freeze exposed skin in seconds; blizzards can reduce
visibility to a few feet, months of darkness and seemingly endless expanses of snow and ice.
Most of Antarctica is covered with vast areas of snow and ice, which reflect about 75% of the
incoming solar radiation.
summer season when Antarctica has continues day for few months that causes ionization at a
constant rate in the upper atmosphere due to solar radiations and excessive heat. A regular
measurement of geo-magnetic activities at the permanent research centers of all the nations is
being carried out. The data records pertaining to this is being properly analyzed and utilized
for navigation purposes. Information can be obtained regarding the movement of magnetic
poles. One can perform a complete analysis related to these phenomenas and it could be
utilized for social welfare.
North and south poles are maintaining the energy balance of the world. The energy from
atmosphere and oceans coming to the poles loses its identity and converts to thermal
radiations. The cold winds in this region combines with hot air of lower latitudes to form
clouds thereby regulating the global environment. To predict the changes in atmosphere in the
future it is necessary to analyze the trend in the past. This continent is ideal for such types of
analysis. The ice that has been accumulating from years withholds natural secrets amongst its
layers. Antarctica is a Scientists Paradise. Science is the Principle human activity in
Antarctica. It is a pristine laboratory and its ice core archives climate history of the past. The
unique nature of the region provides a living laboratory where scientists can measure the
effects of changes in the environment. Ongoing research is crucial to the understanding and
monitoring of the global warming, ozone depletion and atmospheric pollution.
The important research work associated with this place is the discovery of ozone hole,
gradual increase in temperature of Earth, information regarding presence of antifreeze
chemical in the body of fishes. Atmospheric research is one of the areas for which Antarctica
provides a rich ground. The fundamental knowledge gained here is currently being used for
practical applications and has potential promise for future applications as well. Antarctic
ecosystems are ideal for biological research due to many factors. Few life forms survive
above the ice because of harsh environmental conditions. A simple, land-based ecosystem is
easier to study here. Thick Antarctic ice sheets provide one of the best records for past
climate change. Ice cores can reveal patterns of mean air temperature, evidence of major
volcanic eruptions and composition of the atmosphere. Antarctica and the surrounding areas
are natural laboratories for scientific research that cannot be done anywhere else on Earth.
Geophysicists can take advantage of Antarctica's ideal conditions to study the effects of solar
radiation on the Earth's magnetic field.
The geographical location of Antarctica has been a field of special interest for the Indian
scientists. Pacific and Atlantic Ocean meet both the poles of earth but Indian Ocean has land
in the north and Antarctic Ocean in the south. Indian Ocean captures the enormous energy
available here and carries it across to the country. This continent is called as key of the
weather of the world. Information regarding weather, climate and water can be obtained from
this continent. Because of its geographical location its the best and static location for
atmospheric researches. Scientists pertaining to geography and environmental study have
accepted this as their prime research area. This land is primarily useful for researches
pertaining to pure and applied science areas.
ozone levels that were presumed to be erroneous. It was also informed that in 1970 the
thickness of this layer was almost double than is 1981 and the rate of ozone depletion had
picked up pace. The exceptionally cold temperatures in Antarctica play a role in ozone losses.
The cooled air gets settled on the poles because of which chlorine gets accumulated on
troposphere.
During summer season at Antarctica the sun is present for all 24 hours. The reaction of
chlorine and ozone continue to occur all the time in presence of sunlight. This depletes the
ozone layer. During September and October the hole widens to cover the whole of Antarctica
and southern Australia. Hole in ozone layer has been observed on North Pole also.
Springtime ozone hole in Antarctica provides favorable atmospheric conditions for observing
the celestial dome. This window enables scientists to probe the structure of the Sun and the
universe with novel precision.
Evidence for levels of global pollution by industry, agriculture and transport sectors is
frozen into the Antarctic ice. Antarctic 4.75 km thick ice sheet is a record of past climate for
the last 5,00,000 years. Antarctica is a vital area of attraction not only due to its scientific
value but also due to its geopolitical and commercial values. The winter season here is from
April to September and summers are from October to March. In the winter season this
continent doubles itself as the ocean around these freezes. As it occupies positions on the pole
hence it has nearly six months of darkness and six months of day. Indian station Maitri
located on Shrimachar Mountain has day from 22nd November to 24th January and continuous
night from 31st May to 20th July. This continent is hit by most severe adversity of climate. The
minimum temperature recorded here is -89.2C at the Vostok centre of Russia in 1983. The
wind velocity and velocity of storms recorded here has been 320 km/h. The windstorms and
hailstorms continue for days together. Calamities like breaking of ice, breaking of ice shelf,
formation of icebergs and its flow into the waters for Kms together, formations of rifts in ice,
render an utmost challenging and almost impossible working environment to people engaged
in researches here. The reflection of sunrays due to ice almost renders the researchers to
continue their work and increases the chances of getting lost.
Antarctic can be compared to a desert also. Any area of land that has less than 254 mm
(10 inches) rain than it is called as desert. There is almost no rainfall here even if rain falls it
is in the form of snow. Hence its called as a cold desert. During winter season the area of sea
ice increases to 20 million sq. kms, which is larger than the total area of the continent. Every
year almost 85% of this part melts in the summer seasons.
The whole game is of the weather; even a smallest flaw in understanding the climate can
turn disastrous. There must be a thousand mysteries that lie unsolved in Antarctica. The more
mysteries that are unfolded the more hidden mysteries line up. There are countless mysteries
in Antarctica being almost impossible to count.
It is strange but a fact that Antarctica plays a vital role in the weather and climate
mechanism of the whole earth. A key role is played by Polar Regions in regulating the
climate of the world. To derive facts and figures related to such issues an in-depth analysis of
the climatic mechanism at Antarctica has to be done. About 97.6% of ice of this continent
inclusive of the oceanic ice is very sensitive towards the global warming. The ice holds
enormous reserve of water inside it. The risk of increase in sea level may arise by melting of
this ice. The whole coastal region is surrounded by winds hence sudden changes of weather
are natural. The climate of this place is considered to be the most hazardous amongst all
continents. The strong winds due to gravitation give birth to storms, which flow from lower
regions of the continent to upper regions. Keeping the diversity and importance of the
weather and climate of this region January Ist, 1957-58 was celebrated as International
geophysical year. Along with these celebrations all the nations started the race of exploration
of this quiet Zone. A non-governmental organization named SCAR (Scientific committee on
Antarctic Research) was set up in 1958. The committee acts as a regulator for nations doing
research here and also establishes co-operation between these nations. A new avenue was
opened in the field of Metrology in Antarctica after establishment of IGY and SCAR.
Antarctica has day for 24 hours in summer and 24 hours of night in winter. We know that
the earth revolves around the sun and simultaneously rotates on its axis also. The earth is
tilted 23 on its axis. During revolution half of the earth that faces sun has day and the
other half portion that lies in the dark has night. Antarctica lies on the south pole of the Earth
and when there is day on the North Pole the South Pole has complete night.
The Earths South and North Pole plays an important role in research of global weather
and climate because heat from lower latitudes is transported to high latitudes through
meteorological process and is than radiated from polar latitudes at larger wavelengths.
Analysis of meteorological datas collected here is of paramount importance in modeling of
global weather and climate. The weather of Antarctica is made up of uncertainties, nonconsistencies and hindrances. There is no similarity and balance in the climatic factors here.
Many nations have set up static or permanents centers in the white continent and have
established their laboratories regarding weather and climate. The environmentalists here
conduct research on weather related factors of this place. Antarctica affects the atmospheric
and oceanic cycle inside the southern hemisphere hence has an impact on the atmosphere and
weather of the world. The ice here acts as a heat absorber, thereby regulates the sea level. To
know the environment of the world completely its essential to know about energy balance of
this white continent. This frozen ice has detained mysteries of climate of past several
thousands year.
To calculate weather of a particular place the data of surroundings is essential. To predict
weather of an area the data both of the surface and upper atmosphere of various regions is
required. There are huge uninhabited areas as well as a major area is occupied by water hence
the complete zone cannot be inspected and no related datas can be obtained. The various
weather satellites set up by several nations have achieved laurels in the area of weather
predictions. To give exact prediction of weather this place is only partially suitable. The
climate here depends on the weather of the sea, which is devoid of any static laboratories.
The ice of Antarctica is a key factor of the weather mechanism of earth. The whole
continent is surrounded by ice, which spreads to 18 million sq. kms during winters and melts
to two sq. kms during the summer season. The ice is quiet sensitive to these changes of
weather. To collect figures and samples from this ice covered regions is difficult and
hazardous but has been eased by remote sensing.
From time to time scientists were able to extract huge ice cores to analyze the life
structures and climate of the past. This would help in knowing the changes of the atmosphere
that has undergone in the past. This information is related with changes in temperature,
carbon dioxide, ozone level, atmospheric pressure etc. Low pressure group of air move
southward from 500 to 700 along with regular wind. At times its a group of 5 to 6 slabs at low
pressure which have impact on surrounding of Antarctica move eastward at a speed of 90
km/hour. Cold winds towards the coastal region gives rise to high velocity snowstorms,
which may occur for a few hours or several moths altogether.
In the early years information was collected to derive normal conditions but now
permanent laboratories have been constructed to assist research. Research using hi-tech
devices was initiated only in the sixties. International Antarctic Analysis centre was set up to
analyze the data available with all the nations collected from work of several years. The data
held was also published so that it could be used to unravel mysteries of weather. Later use of
satellite was made to predict weather, climate, pressure, temperature, humidity, direction and
velocity of wind etc. Correct information of all these factors was available only because of the
satellites used. After the use of these technologies it became easy to know about the location
of low-pressure area.
To know the climatic hazards of this region information for research is being made
available through world climate research program and World Meteorological Organization.
Today efforts are being made to know the degree to which weather can be predicted and the
human activities affects the weather. Some scientists belonging to member nations of SCAR
are conducting research pertaining to movement of ice, its thickness, energy flux and
activities in the atmosphere.
The layer of ice serves as a mirror to solar radiations coming from the top and reflects
these rays back into the atmosphere. There by this continent shreds energy in the form of
radiations and acts like a global energy sink affects the weather. To know in detail about the
weather and climate total devotion and carefulness in required. It is essential requirement to
analyze the surroundings of Antarctica and the ocean and also to analyze the reactivity and
effect on oceanic waters. Weather experts in various centers have analyzed pieces of ice to
know chemical reactions along with freezing and changes in atmosphere several decades
before. Russian scientists have analyzed the freezing rate, atmospheric temperature of
oxygen, percentage of compounds of hydrocarbon in an ice slab extracted from 2000 mts
under ice. With more human intervention for research the carbon dioxide percentage has
increased causing an increase in the green house effect due to which an increase in the
warmth of the continent is observed. On analysis of pieces of ice from the layer for the gases
enclosed a rise in percentage of CO2 has been proved. Two hundred years ago the percentage
of CO2 was 200 ppm, which went upto 280 ppm hundred years ago and now it has risen to
340 ppm. Many years ago the reason for low CO2 percentage was dissolution of CO2 in cold
water, which has better dissolution capacity than hot water. Thus the glaciers were more static
pertaining to low CO2.
In the recent years almost all nations have established their automatic weather stations in
this region. Implementing the modern hi-tech devices information regarding Antarctic and the
rest of the world can be predicted at least a week before. This white continent has become an
inseparable part of the climate and weather mechanism of the globe. It is even possible to
calculate the weather of high latitudes easily.
From the first expedition, India had started gathering information related to weather for
research purposes. Regular per minute inspection of the velocity of air, state of clouds,
visibility, humidity, vapour etc. by the Indian metrology department is going on. The
10
laboratories have all self-start and automatic devices. These laboratories experimenting on
diffused solar radiations, surface ozone, total ozone, ultra violet radiation, measurement of
Green House Gases etc. The prior prediction of weather is useful especially for scientists who
perform research work in the Antarctic fields.
The average temperature of Antarctica is - 490C. On the southern pole the sun shines for
24 hours and for nearly 6 months but if one feels that a little warmth can be felt due to sun it
is wrong. The outside world avails information regarding Antarctic from datas available
through satellite. Satellite can be used to obtain information regarding temperature of water,
ice layer on water, construction and expansion of ozone layer.
The recording of the climate of this region started in 1950. Through the data and pictures
available from satellite the state of clouds, velocity and movement of storms, formation of ice
and its distribution and various environmental processes can be updated. Various global
climatic models clearly indicate that there would be enormous changes in the climate of
Antarctica during the next 100 years which would cause an increase in temperature that
would in turn increase the snow melt. The available climatic models have failed to produce
reliable data of the changes in weather hence more research is required in this field.
Several experiments have proved that there are certain changes occurring in the upper
atmosphere. In ionosphere in the F- region (300 kms) the peak of concentration of electron
density from the past 38 years has come down by 8 kms. In the lower atmosphere due to
increase in concentration of green house gases the temperature has increased but the upper
atmosphere has become relatively colder. The various theoretical analyses have shown that
the layers have decreased. Southern ocean acts like a sink in global carbon cycle. The layers
of ice, oceanic ice and oceans are active factors of climate.
The polar plates being formed over Antarctica render it coolest. There is extreme cold
due to flow of cold air. The air flows from cold mountains at a height of 9000 feet downwards
and flows towards the coastal region. The rotation of earth deflects the air to left side.
Researchers believe that these cold winds have major effects up to long distances. In the
coastal areas the wind velocity is 150 meters per hour.
The 5 meter thick ice of this continent deposited from 5 lakhs years encloses the climate
of the past. These gas bubbles enclosed in the ice are a witness of pollution caused by
environmental gases, industrialization, atomic bomb and fertilizers used for crops. The future
climate and weather can be predicted on the basis of research on theses factors. Every year
during winters about 7 million sq. kms of ice deposits around Antarctica, which melts during
summers. This is the biggest event associated with weather. Scientists have discovered two
more sub glacial lakes, raising renewed speculation of the existence of living ecosystems
below the ice. The one lake is called "90E" because it lies along that longitude and is now
the second biggest sub glacial lake in Antarctica, while the other is named after the Russian
station above the lake called Sovetskaya. These lakes were known of before but this is the
first time that scientists have been able to assess their size using satellites and ground
penetrating radar equipment.
Scientists and experts coming to Antarctica are conducting research on various problems.
Zoologists are analyzing the behavior of animals residing here and are trying to find out
reasons due to which they can survive in this hostile environment. Some scientist has
discovered very small plants growing inside rocks. The biologists analyzes water of lakes
covered to find out fecal coli form and other chemical contaminations. Environmental and
weather experts analyze the climate of Antarctica and trying to discover reasons of the
11
12
provide a constituted outlook to the research happening here so that it could benefit all
nations.
REFERENCES
Chaturvedi, A:. Rochak avam Romanchak Antarctica: Prabhat Prakashan, Delhi, pp.1-256,
ISBN No.81-7315-532-1, (2006).
Khare, N.: Dharkta Mahadeep, NCAOR, Goa (2000).
Girija R.:-Atmospheric research from Antarctica, the Indian contribution, pp. 1-104, IIG
Bombay (1993).
Purohit, P.K.: Antarctic: Vigyan Avam Chunotion., Madhya Pradesh Hindi Granth Acadamy
Bhopal, pp. 1-116,(2004).
Purohit, P.K.: Antarctic: Aakarshan Kyon? Indira Publishing House Bhopal, pp.1-175, ISSN
No.818910703-8, (2005).
Purohit, P.K.: Antarctic: Where the Silence Speaks, Indira Publishing House Bhopal, pp. 1180, ISSN No. 8189107-14-3 (2006).
Chapter 2
E-mail: hndutta@gmail.com
Roorkee Engineering and Management Technology Institute,
Shamli-247774, U.P. India
2
Pawan K. Sharma, Department of Chemistry, Kurukshetra University,
Kurukshetra-132 119, India
3
N.C. Deb, Indian Statistical Institute, 203 B.T. Road, Kolkata-700 108, India
4
Laxmi Bishnoi, Government Girl's P.G.College, Gurgaon-122 001, India
1
14
1. INTRODUCTION
Antarctica is a magnificent display of interaction between air and the various phases of
water (vapor, liquid and solid) in a pristine atmosphere, where the direct solar energy reaches
the Pole for six months of the year at very low elevation angle, and for the other six months of
the year, it is dark. This has already led to the formation of a dome shaped, permanent ice cap,
which is about 4 km thick in the interior of the continent, having an area of about 14.5 m km2
(or a circle with a radius of about 2000 km).
Antarctica is surrounded from all around by an equally magnificent ocean called as the
Southern Ocean and both the Antarctic continent and its surrounding ocean are connected
through the most efficiently coupled ocean-ice-air system (Bailey, 2000; Schellenberg et al.,
2002; Liu et al., 2004; Hall and Visbeck, 2002; Parkinson, 2004; Simmonds and King, 2004;
Naithani, 1995). The Southern Ocean is dominated by a yearly variation of sea ice; its
maximum area is about 19 million km2 in late winter and the minimum about 4 million km2
during late summer (Arrigo and Thomas, 2004). The moment sea ice forms, it reflects the
solar radiation, thus enhances the sea ice growth. Also, being a poor conductor of heat (Van
den Broeke et al., 2006; Lefebvre and Goosse, 2008), it separates the warm ocean water and
the colder atmosphere and thus stops the transfer of heat from ocean to atmosphere and vice
versa. Moreover, when sea ice forms, brine is rejected into the underlying ocean creating a
layer of saline, dense water decreasing the vertical stability at the base of the oceanic mixed
layer (Kwok and Comiso, 2002). Subsequently, entrainment causes a negative feedback
between the ocean and sea ice, whereby the deeper, warmer ocean water is brought to the
surface, increasing oceanic heat flux and inhibiting sea ice growth (Martinson, 1990;
Marsland and Wolff, 1998). The heat flux is greatest initially when brine rejected by the
rapidly growing ice sets up thermohaline convection in the underlying water. As the rate of
ice growth and brine rejection decreases, so does the convection and the heat flux to the lower
ice boundary drops off.
During summer, sea ice melts mostly by warming of the ocean mixed layer through heat
input (mainly solar radiation) in open water areas (Ohshima and Nihashi, 2005). Also, a large
number of low pressure areas in summer result ocean swells and high winds breaking the ice
into large pieces that move under the influence of wind and currents. (Fast-ice is sea-ice that
is held fast to the continent.) Pack ice can change in a matter of hours from being open and
navigable to densely packed and impassible.
The large seasonal fluctuation of the sea ice cover affects the exchange of energy, mass
and momentum between the ocean and atmosphere, and is extremely important in controlling
the existence of Antarctic continent and its interaction with the whole globe (Lefebvre and
Goosse, 2005, 2008; Simmonds, 2003; Thattermann and Levermann, 2009; Busalacchi, 2004;
Meskhidze and Nenes, 2006; Ito et al., 2010; Ved Parkash, 2008; Liss et al., 2004).
The Southern ocean's circulation system has another unique feature known as the
Antarctic Circumpolar Current (ACC), which is both the longest and the strongest current in
the ocean carrying a volume transport of 130Sv (1 Sverdrup=1Sv=1106m3s1) along a
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
15
24000 km path encircling Antarctica (Gille, 1994). The ACC is also unique because no
continental barriers exist in the latitudes spanning Drake Passage (the gap between South
America and the Antarctic Peninsula), which allows the current to close upon itself in a
circumpolar loop and is the primary means of inter-basin exchange of heat, carbon dioxide,
chemicals, biology and other tracers. Despite its essential role in the climate system, the ACC
remains one of the most poorly understood currents in the ocean (Thompson, 2008). At the
same time, deep waters in the Southern Ocean are rich in dissolved inorganic carbon (DIC)
and depleted in oxygen. When circulation brings these waters into the mixed layer, the
soluble gases are exchanged at the air-sea interface. The formation of deep mixed layers
combined with high biological productivity make the Southern Hemisphere extra-tropical
oceans an important component of the global carbon cycle (Bishop and Wood, 2009).
16
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
17
cleaner, as the suspended particulate matter in the air is brought down by the falling
snow flakes and buried in the ice (Wang et al., 2008; Boucher et al., 2003; Eisele et
al., 2008; Stohl and Sodemann, 2010; Gajananda et al., 2004, 2007). This cleaning of
the atmosphere is essential for the radiative part to be efficiently active over the
entire Southern Ocean. The cleaning of the atmosphere also happens even during
summer, when the cyclones lead to snowfall. This is an absolutely essential
component of the entire Southern Ocean environment as for the formation of the sea
ice, it is necessary that the radiative energy must escape to sky without any hindrance
or modification etc. (Chamberlain et al., 2000; Lawrence et al., 2004; Naithani et al.,
1994).
In summer, cyclones may also result in some rainfall events at the periphery of the
continent (Deb et al., 1999; Ved Parkash, 2008). In fact, over the past thousands of years, the
loss of Antarctic ice had been lower than the total amount it gained and thats how, Antarctica
is holding its physical structure and is participating in the controlling the global temperature
and many other important features.
Even in the past, there has been no change in the snowfall over the continent
since the IGY (Monaghan et al., 2006) and Antarctica seems to be stable in some respect
(Gudmundsson and Jenkins, 2009). However, many signals or worries are related to the
global warming, which makes the subject of ocean-ice-air interaction to be an important
science subject (van de Berg et al., 2006; Justino et al., 2010; Peck, 2005; van den Broeke,
2008; Bindschadler, 2006; Steig et al., 2009; Gille, 2008; Zazulie et al., 2010; Atkinson et al.,
2004). In fact, to strengthen our knowledge about various aspects of Antarctica, many
programs are in progress (Rignot, 2002; Frey et al., 2009).
Once the katabatic winds are forced to blow over the ocean, they move up the relatively
warm and humid air. As this humid air moves up, it moves into a region of low pressure and
cools. The associated water vapor also cools to form water droplets or directly into snow,
releasing latent heat. This release of latent heat creates a low pressure area, with the result;
more air is sucked into this region. Thus, it triggers the formation of a well defined or well
developed cyclone over the ocean, which has to move towards the continent.
The cyclonic air from the ocean side is relatively much warmer than the existing
Antarctic air, leading to warming of the atmosphere, which is more noticeable in winter
months (Naithani, 1995; Gajananda, 2002). In fact, there are events over Antarctica, wherein
a widespread warming has been observed both during local summer and winter seasons
(Enomoto et al., 1998; Deb et al., 1999; Van As et al., 2007).
In summer, the katabatic winds are weaker but the number of lows per month is higher in
summer due to advection of water vapor (Naithani, 1995; Kumar, 2002; Carrasco, 2003; Ved
Parkash, 2008). At the same time, the invaded moisture by the cyclones may lead to the
formation of fog over the periphery of the continent (Gera et al., 2002; Walden et al., 2003;
Gajananda et al., 2003; Gajananada et al., 2007).
These cyclones approaching from lower latitudes (moving from west to east), become the
main sources of heat for the atmosphere, and the atmosphere-surface heat transfer takes place
through turbulent mixing and longwave radiation, the latter is dominated by clouds. The
cyclones are also responsible for warming of planetary boundary layer around the periphery
of the continent during winter.
18
Also, the cyclones replenish the loss of ice in the form of fresh snowfall all over the
continent and around the ocean. The freezing ice in winter and the falling snow cover almost
all the exposed parts even at the periphery of Antarctica, thus prepare the entire continent to
be ready to reflect the falling Suns energy during the local summer.
The atmospheric research in Antarctica is increasingly important as many processes
taking place in the atmosphere are still barely understood (Anisimov et al., 2007). This is
particularly true for exchange of gases and particles between ocean and atmosphere, an area
of study which is currently receiving much attention (Meskhidze and Nenes, 2006; Zorn et
al., 2008).
Since the Antarctic atmosphere is absolutely clear, aerosol particles play an important
role in the atmosphere because of their effects on the radiation budget and consequently on
climate and climate change, which is in particular true for aerosols from marine environments
(ODowd et al., 2007). Currently, the focus is on their formation and understanding which
gaseous compounds participate in it (ODowd et al., 2007) and the atmospheric processes
over the most complex situation caused by the water, ice and air (Andreas et al., 2004;
Bishnoi et al., 2005; Dutta et al., 2004 ).
In the present Chapter, we shall concentrate only on the atmospheric phenomena, which
have been captured by a shipborne acoustic sounder deployed onboard the ship Megdalena
Oldendroff, which sailed along the periphery of the east Antarctic ocean in the year 2002.
3. SODAR MEASUREMENTS
The group at the National Physical Laboratory, New Delhi had established an acoustic
sounder (sodar) at the Indian Antarctic station, Maitri (70.76o S; 11.73o E; elevation 117m) in
the year 1990-91 (Dutta et al., 1991; Dutta and Naithani, 1994; Naithani and Dutta, 1995;
Gajananda et al., 2004, 2007; Kumar et al., 2007; Gera et al., 2010). This development had
made India as the sixth country in the world to have a boundary layer program over
Antarctica and since then, attempts have been made to install a sodar system onboard the
ships sailing to Antarctica as part of various Indian Antarctic expeditions (Dutta et al., 1993,
1999, 2004, 2007; Bishnoi et al., 2005).
The most successful attempt for shipborne operation was made during the 21st Indian
Scientific Expedition in the year 2002, when the system was installed onboard the ship
Megdalena Oldendroff, which sailed from South Africa to Antarctica. The team established it
successfully and operated PC controlled system as shown in Figure 1. Figure 1 shows
photograph of the antenna mounted onboard the ship and that of Dr Pawan Kurmar Sharma
who installed and operated the system successfully.
The system was placed on the ship around January 19, 2002 and after making several
checks, it finally started functioning from January 22, 2002. At the same time, the ship was
sailing; its pathway is shown in Figure 2. The ship actually sailed to the German Antarctic
station, Neumayer, which is situated on the shelf itself. It is important to note that the ship
cannot be anchored along the shelf during the cyclonic conditions for various reasons and
therefore it is kept in motion. The square shown in Figure 2. indicates that the ship remained
in this area from February 3 to March 11, 2002.
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
19
Figure 1. Shipborne acoustic sounder antenna onboard the ship Megdalena Oldendroff and Dr Pawan
Kumar Sharma operating the system electronics.
Figure 2. It indicates latitude and longitude that ship followed during its journey in Antarctica. The box
indicates that the ship remained in this area from February 3 to March 11, 2002.
20
Out of this useful data of 476 hours, 69.9% of the time, it was surface based inversion,
followed by thermal convection for 15.3% of the time and elevate inversion for 14.7% of the
time (Figure 3). This indicates that even during sunny hours, it is the stability in the lower
atmosphere, which dominates the lower marine planetary boundary layer close to the
periphery of the east Antarctic continent.
Thermal
convection
15.3%
Elevated
inversion
14.7%
Surface based inversion
69.9%
Figure 3. Out of the total 476 hours of noise free recording by the shipborne acoustic sounder, surface
based inversions dominated the occurrence followed by elevated layers and the thermal convection.
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
21
lines, indicating wind noise. The surface based inversion thickness is an important parameter
and is plotted in Figure 5. This shows that surface based inversions have a high variability in
terms of thickness, which can as low as just 50 m or as high as 325m. Of course, the most
probable thickness lies around 150 m. However, it is important to note that the data given in
this figure cannot be taken for any strong or definite conclusion, these are just preliminary
observations actually meant to show that the shipborne acoustic sounder can function
normally even under the most adverse Antarctic conditions, recording variability in the
thermal structures of the lowest PBL.
Figure 4. An example of a surface based inversion recorded by a shipborne Acoustic sounder observed
over the southern ocean along the eastern coast of Antarctica. This echogram directly gives the
thickness of inversion and is an important parameter in many atmospheric applications.
Figure 5. Statistical distribution of thickness of surface based inversion in various categories. The SBI
thickness has a high variability but is most probable around 150 m.
22
Figure 6. Thermal plumes are an important atmospheric phenomenon in the Antarctic atmosphere as
they represent surface air to be much warmer than the air aloft. The warmer surface air rushes up to
form an inverted cone on the acoustic sounder echograms. The height up to which these thermal go
represents the thermal mixing in the atmosphere.
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
23
Figure 7. The occurrence of thermal convection in various height ranges as recorded by the shipborne
acoustic sounder over the east Antarctic coast. As expected, it has a high variability but the most
probable height attained by the plumes is around 300 m. This is an important finding and shall be useful
in many models.
Figure 8. Elevated inversions in the Antarctic environment are an expected phenomenon in the PBL
Shipborne acoustic sounder has revealed a variety of these layers, which have shown a high degree of
variability in the height but thickness is generally between 50-80 m.
24
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
25
Figure 9. Highly variable facsimile picture shows a turbulent planetary boundary layer over the
Southern Ocean.
Figure 10. Dominance of surface based inversion is the key for the survival of Antarctica but patches of
thermal convection are indicators of atmospheric variability over the ocean.
26
Figure 11 a. An extremely important and unusual event of continuous thermal convection over the east
Antarctic ocean is a subject of great scientific value and needs in-depth investigations.
This is something unique as if we see the sunlight hours, the convection can only be
expected during the peak sunlight hours and that too, if we presume that there was no mixing
over the ocean. On the other hand, if the oceanic water is warmer than the air, then the
thermal convection can be sustained on a prolonged basis. In any case, thermal convection
indicates that the surface is at least 3-4oC warmer than the air aloft and the horizontal mixing
is low. It is the time when surface of ocean transports its heat to the atmosphere.
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
27
Figure 11.b. Dull thermal convection continuing from the previous day, is again an extremely important
phenomenon and calls of well planned PBL studies over the east Antarctic Ocean, which is holding
many natural secrets. The most important part is that even under high winds beyond 1030 hrs., dull
convection continued till 2115 hrs.
6.1.1. Discussion
It is important to note that there was a surprising event over the east Antarctic
region in early 2002, when the break-up of the Larsen B ice shelf was observed
(http://www.coolantarctica.com/Antarctica%20fact%20file/science/global_warming.htm).
This event has been attributed to the effects of global warming. That it occurred is beyond
dispute and that it is a result of the warming of the Antarctic Peninsula where it is situated is
also beyond dispute. What remains unclear is whether or not this is a taste of things to come
and an indicator of an Antarctic-wide phenomena or simply a localized result of the localized
warming of the Antarctic Peninsula region alone.
The Larsen B ice shelf was about 220m thick (720 feet) and during a 35 day period in
early 2002 lost about 3,250 km2 of ice into the ocean. It is thought to have been in existence
for at least 400 years prior to this and probably as long as 12,000 years since the end of the
last ice age.
Such disintegration in such a short time period is therefore an extremely significant event.
What now remains of the Larsen B is about 40% of what was there in 1995. It had been
breaking up at what was considered to be a rapid rate anyway before this major event. The
break-up is thought to be a consequence of higher temperatures and large amounts of summer
melt-water running down crevasses in the ice shelf so speeding the disintegration process.
28
Overall in the Antarctic Peninsula, seven ice shelves have between them declined in area
by about 13,500 km2 since 1974. This melting on such a large scale cannot be explained just
on the basis of warming of the air, it has to be basically warming of the water, which had
dissolved the whole mass of ice. This warming or warm water must have been below the ship
on February 2-3, leading to the formation of thermal convection observed for over 24 hours.
A series of Southern Hemisphere experiments have been performed to study turbulent
convection on a continental shelfslope placed in a large rotating tank filled with fresh water
(Liang et al., 2008).
The upward ocean heat transport is generated by oceanic advection, diffusion, and
convective overturning. In the Southern Ocean, convection occurs along the continental
shelves of Antarctica as well as in the open ocean owing to ice formation and associated salt
rejection (Baines and Condie 1998). Bitz et al. (2006) have demonstrated the strong influence
of sea ice on convection and the upward ocean heat transport through freshwater transport,
which makes the surface waters more stable in a greenhouse warming scenario.
In support of the thermal convection, it is important to note that the synoptic pictures
show that there was certainly warming over the investigation area (Figure 12).
Figure 12. Rising layer is a common phenomenon over the land and is well understood. But, over ocean
and that too in Antarctica, where the ocean is the most turbulent and winds are strong, this is something
unique and needs in-depth investigation.
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
29
Figure 13. Rising layer over Antarctic ocean is a unique feature on acoustic sounding as it hard to
expect that the oceanic/ icy surface will not transfer energy to its own medium, rather only the upper
surface will get warmer to create convection and sustain it for at least 6 hours.
In fact, it is the thermal plumes, which lift the inversion at the same time, erode it from
the lower side of the inversion. In the case of brisk heating, the plumes may penetrate into the
inversion and may break it within a short span of time. On the other hand, like in winter, it
may take several hours or the upward moving inversion may not break at all, it may remain as
a suspended layer throughout the day and then return back to ground in the evening as the
heating subsides (Dutta et al., 1994; Choudhury and Mitra, 2004; Kumar et al., 2010).
Over the Indian Antarctic station Maitri, an acoustic sounder was deployed in the year
1990 and here also, over a period of six years, some incidences of rising layer were noticed
but such cases were not very prominent (Naithani, 1995).
On the other hand, on the ship, a case of rising layer was observed and it very unique as
over ocean, it cannot be expected that either ocean or ice will behave like Earth. In the ocean,
some amount of mixing will always be expected and over the ice, the melted water under
solar heating shall always dissolve more ice. But under exceptional circumstances, it is just
possible that the energy exchange underneath the water or ice gets limited and the energy
exchange takes place between the wet /icy surface and the air. But, it would certainly require
a high pressure area over the observational site to ensure calm weather (calm ocean) and low
winds.
6.2.1. Discussion
We have scanned the whole literature to get a support for our observation but there is
nothing that can be quoted with firmness. However, it is important to note that for a rising
layer, the heating has to be continuous on the ground to support the development of thermal
plumes underneath the rising layer. This will only be available if the mslp is high so that the
atmosphere is clear to form the steep inversion and in the morning to receive the heat to warm
up or erode the inversion. The atmosphere, of course, has to be still.
30
Figure 14. Isobaric contours of surface msl recorded on January 22, 2002 indicate high Pressure zone
(marked as H) around the ship position.
Figure 15. The sea ice erosion along the eastern coast of Antarctica has been severe in the months of
January-March, 2002. Sreenivasan and Majumdar (2006).
This support is available for the synoptic pictures as seen in Figure 14. It is clear from
this figure that there was indeed a high pressure area or zone between two consecutive
cyclones. This is what normally happens in Antarctica (Naithani, 1995; Kumar, 2002;
Gajananda et al., 2004; Ved Parkash, 2008) and the weather is altogether different in two
regimes (the low pressure area and the high pressure area).
The other support is from the surface temperature (Figure 15), which shows the warming
over the eastern zone of Antarctica and relatively open ocean so that its albedo is low.
An Insight into the Ocean-Ice-Air Interactions over the East Antarctic Marine
31
Actually, the whole month of January was getting warmer as is seen from the published
work of Sreenivasan and Majumdar (2006), wherein authors have reported the unusual
melting and the figures are presented here as Figure 15. The calculated estimates are given in
Table 1.
Table 1. Sea ice areal extents and depletion statistics
Date
15.1.2002
23.1.2002
Weekly depletion
Million km2
5.023
4.348
1.160
0.675
CONCLUSION
Antarctica is the least explored and still a mysteries continent on Earth; through acoustic
sounding of the atmosphere, we have explored two of its unique secrets for the first time in
the history of mankind. Both of these events throw a light on the need to continue acoustic
sounding program with better coordination and planning for giving a proper interpretation for
the better understanding of the water-ice-air-radiation interactions over Antarctica. After all,
Antarctica holds the largest fresh water stock on Earth, the key to absorb CO2, millions of
unique species and much more for the betterment of mankind on Earth.
ACKNOWLEDGMENTS
The authors are grateful to Dr. B.S. Gera, who supported the development of shipborne
acoustic sounder in India. Authors would like to thank members of various Indian Scientific
Expedition teams, who helped in the installation of acoustic sounders onboard various ships
sailing over east Antarctica. Thanks are also due to the Chairman, CSIR-SCAR for selecting
and providing field support to various NPL Antarctic teams.
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36
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Chapter 3
LAND-ICE-AIR-OCEAN INTERACTIONS
IN THE SCHIRMACHER OASIS, EAST ANTARCTICA
Khwairakpam Gajananda1 *, H. N. Dutta2 and Victor E Lagun3
ABSTRACT
Study of the land-ice-air-ocean interaction over the Schirmacher Oasis (SO) of east
Antarctica was carried out during the years 1999-2000. The research revealed that a
unique ecosystem prevails over this oasis, where all the physical and biological
components interact in a complex pattern. SO ecosystem is heterogeneous in nature and
the estimated biomass production is low at a value of 22.5 gm m-2. The lakes are
oligotrophic in nature and are of fresh water lakes. Food chain of SO is very simple and
short and the energy cycle is poor due to less sunlight during austral winter. The diversity
of the flora and fauna is poor and is dominated by poikilohydric microorganisms. About
34 species of primitive flora were observed and only 6 primitive invertebrate fauna were
recorded. Only four species of birds were observed in SO. The migratory birds from subAntarctic islands may introduce some non-indigenous forms of plants, animals and
microorganisms species to SO, but only cold tolerant variety of organisms survived.
Average organic carbon content of the oasis is 1.58%, which is poor in comparison to
other ecosystem of the world. The microbial enzyme activity or dehydrogenase activity is
0.008 mg TPF g soil-1 day-1 suggesting a very less microbial activity for substrate
decomposition. Experimental investigation suggested that the climate at Maitri is
dominated by the extreme contrasts between the seasonal inputs of solar radiation. SO
experiences sub-zero mean temperature throughout the year except in the peak summer
months (December and January). Pressure (average 986.5 mb) forms half yearly cycle
and influenced for the formation of cyclones. The humidity and precipitation are low but
have significant relationship for the growth of microorganisms. Convective atmospheric
phenomena during austral summer have the potential for dispersal of the microorganisms
*
40
1. INTRODUCTION
Antarctica presents the most efficient and the most interactive land-ice-air-ocean system
in the world (Dutta et al., 2007). The land-ice-air-ocean interactive system supports extreme
climatic conditions in the interior of the continent. With the result, only micro flora and fauna
have been able to evolve with time and survive, forming one of the most important subjects in
Antarctica for human understanding (Gajananda 2003).
In Antarctica, only Oasis regions get deglaciated during local summer and therefore,
present the most favourable conditions for the survival, growth and function of the micro
flora and fauna in the otherwise harsh climatic conditions. The deglaciation of the east
Antarctic oasis might have started very rapidly, close to the Pleistocene/Holocene boundary,
probably favoured by marine transgression, sea level falls and climatic warming (Melles et
al., 1997). Antarctic continental soils are arid, saline and lacking in organic matter, whereas
maritime soils, in wetter environment, range from structure-less lithosols to frozen peat (Hall
and Walton, 1992). Two important factors in the development and diversity of terrestrial
communities are water availability and the period of exposure since deglaciation.
The retreat of ice sheets offers new sites for colonization by microbes, plants and
animals. The interactions between snow line, freeze-thaw cycles, wet-dry cycles and the
length of the summer are considered as critical in determining the extent and rate of localized
changes in weathering and pedogenesis (Gajananda 2007). The implications of higher
temperatures and differing precipitation regimes are considered in relation to weathering, soil
development and the establishment and development of terrestrial communities of flora and
fauna. A study on this subject will provide a good model of how present soils and
communities developed at the end of the last glacial age. The east Antarctic region, has
relatively young terrestrial and inland water ecosystems, dominated by a few or single
species. These ecosystems offer a great deal of information about species adaptation and
reproduction. Moreover, the coastal region is ideal for the study of dispersal and colonization
across great expanses of ice and ocean (Gajananda 2003). The 8 major coastal oases of
Antarctica provide the migration and immigration of organisms during austral summer. The
migratory birds and Penguins migrate from various sub-Antarctic islands to these regions in
summer, bringing with them various forms of seeds, propagules of plants, spores and minute
invertebrates. The atmospheric boundary layer dynamics over Antarctica has been studied at
many stations by acoustic sounding technique, which provides an in-depth knowledge of the
PBL dynamics (Argentini et al., 1996; Gera et al., 1997; Naithani and Dutta, 1995). Acoustic
sounder finds applications in the areas of fog monitoring (Beran and Hall 1974; Gajananda et
al., 2007). The coastal Antarctic planetary boundary layer (PBL) experiences varying external
influences both from the interior of the continent and due to the moving depressions/cyclones
along the coast (Wendler and Kodama, 1994; Carrasco and Bromwich, 1997). The influence
41
from the interior of the continent is dominated by katabatic flow of winds of varying
intensities, often depositing snow/ice in the form of blowing snow, drift and blizzards (Du
and Bromwich, 1992; Argentini et al., 1996; Braaten, 1997). However, cyclones push
relatively warm and moist air towards the interior of the continent, leading to foggy weather
condition (Naithani and Dutta 1995) and coreless nature of temperature variation over
Antarctica (Wendler and Kodama, 1994; Naithani and Dutta, 1995). In the driest and coldest
habitats, especially where fog and dew are major water sources, desiccation-tolerant algae or
cyanobacteria, bryophytes, lichens may form the only vegetation (Alpert and Oliver 2002). At
the same time, thermal convection over the ocean transports fine living materials and
propagules (Gajananda et al., 2004b). During summer, the atmospheric processes and the
local solar heating make the Antarctic boundary layer to be one of the most dynamical
regions (Kottmeier et al., 1993; Williams and Hacker, 1992 and 1993). It can be mentioned
here that the increasing scientific activities and tourisms over Antarctica has introduced new
and alien species and also unwanted materials are increasing day by day. The extent of its
impact on this pristine environment is yet to be ascertained precisely. Strictly speaking,
vegetation should be considered to be part of the climate system on all timescales. Thus, this
study presents a detailed investigation on almost all aspects of climatic/atmospheric
phenomena in the Schirmacher region that influence both the terrestrial and aquatic ecology
or are directly responsible for ecological control and development of this region. Results
obtained from the analysis of the samples collected over the Schirmacher Oasis (SO
hereafter) during the period December 1999 to January 2000 have been discussed. At the
same time, support of the personal observations related to the number of ecological study and
the simultaneous measurements of the atmospheric parameters, during the above period have
been utilized. Since various ecological factors affect the functioning of life forms in a holistic
manner (i.e. all the factors operate in conjunction and not in isolation), it becomes difficult to
understand the mechanism of the nature of influence by an individual factor. To understand
this mechanism of environmental influences, it is essential to study the effect of each
individual factor separately, thus, by taking into account the concept of an analytical
approach; the following paper has been organized accordingly.
42
interconnect many lakes. The depth of channel entrenchment is different comprising 8-10 m
in the break segments. In mid-summer during the period of intense melting of snowfields and
the glacial slope adjoining the SO, the area of some lakes significantly increases. Numerous
small lakes appear with an area of up to several tens of square meters. By genesis, the lakes of
glacial origin dominate. There are many relict lakes-lagoons located at the boundary between
the Oasis and the ice shelf. Both shallow (3-5 m) and deep lakes (20 to 120 m) are
encountered. Water in the lakes has very low mineral and small hardness. Figure 1. gives the
sampling locations, fresh water lakes, rocky areas, ice shelves, polar icecaps with the two
adjacent east Antarctic researches stations namely Maitri and Novolazarevskaya. SO can be
classified as a true cold desert of Antarctica (Wharton 1993). In this oasis average annual
precipitation (expressed in terms of water) is between 50-150mm (Schwerdtfeger 1979).
Heavy snowfall occurs when cyclonic storms over the surrounding seas push in relatively
warm and moist air over the continent. This moist air freezes and is deposited as snow over
the areas. The region has almost continuous daylight during the local summer and darkness
during the winter. The continental character dominates climate of the SO, except in local
summer when it is predominated by the intensity of solar radiation heating the exposed rocks.
The weather forms depend on the position of the circumpolar trough, solar parameters, flow
of katabatic winds and the position of various cyclones along the periphery of Antarctica. The
circumpolar trough has a biannual movement, leading to the surface pressure level to be the
highest during summer and winter periods. In contrast to the seasons normally observed over
the Northern hemisphere, the seasons of Antarctica have been divided as given in table 1. The
relative air humidity, on an average does not exceed 52% in a year. Under such conditions,
strong evaporation and melting of snow occurs, which is probably, one of the decisive factors
ensuring the existence of the oasis under the current climatic conditions. Table 2. shows the
average climatic characteristics of the SO, calculated from the meteorological data recorded
during 1990-96 at Maitri, Antarctica.
43
Seasons
Summer
2.
Autumn
3.
Winter
4.
Spring
43.9 kcal/cm2
93.8 kcal/cm2
69.6 kcal/cm2
23.9 kcal/cm2
-11C
988.0 mb
10.2 m/s
ESE
52%
309 mm
88 days
0.07 hPa
5.8 points
The group at the National Physical Laboratory (NPL), New Delhi has been primarily
working on the Atmospheric Boundary Layer (ABL) over the SO since 1989-90. During the
years 1990-1997, a monostatic acoustic sounder or Sodar was operated at the Maitri, Station
(Naithani and Dutta 1995). The Sodar measurements indicate stable atmospheric condition
throughout the year (~95%), except in summer, when thermal convection predominates
during the noon hours (~5%). The stable atmosphere is basically due to the extreme
transparency of the atmosphere, leading to the cooling of the icy surfaces. However, heating
of the dry, rocky region, especially in the warmer hours of the day, causes the thermal
convection. In the Antarctic environment, penetration of UV light is much deeper. However,
it is influenced by the influx of suspended particulate material, sea salt etc. by the cyclones
churning and pushing the warm, moist oceanic air towards the continent.
44
45
along the length of SO. The temperature was recorded at the site of collection using simple
thermometer during the sampling session and compared with the daily station temperature
records at Maitri. From all the 14 sites, samples of soil, water (wherever available), rocks and
biological samples were collected in triplicate. The sampling and analysis were carried out
based on the standard methods.
Table 3. Description of the 14 sampling sites at the Schirmacher Oasis
Site No.
1
Habitat type
Swampy lake bank
Lake bank
Site No.
3
Habitat type
Hilly rocky area
Transition of continental
ice and SO rocks
Lichen dominate
7
8
9
10
11
12
13
14
Remark
Little human
activities
Little human
activities
Remark
Little human
activities
Little human
activities
Little human
activities
Little human
activities
Moderate human
activities
High activities of
human
Little human
activities
Little human
activities
Little human
activities
High activities of
human
Little human
activities
Little human
activities
46
Figure 2. Most efficiently coupled Land-Ice-Air-Ocean Interaction System in the World. It maintains
itself strongly but gets influenced by the global changes. (Dutta et al., 2007).
In the process, cyclones churn the ocean to mix the oxygen in the oceanic water,
promoting high rate of biological production. Also, due to the cyclones, long exposure of the
high UV-B doses to the biological organisms is avoided due to creation of turbulent mixing in
the ocean, as well as, due to the development of thick clouds and precipitation. High doses of
UV-B radiations in Antarctica might also be playing a role in degrading the dead organic
matter, thereby releasing various nutrients, which could be readily used by planktons, thereby
rendering Antarctic Ocean as the most productive ocean in the world. At the same time, the
47
cold oceanic water has much higher capacity to retain dissolved oxygen, the most essential
component in the production, growth and function of organisms, as well as to support krills
and various types of fishes in the southern ocean (Dutta et al., 2007). Under natural
conditions, it is found that living organisms are affected by the sum total of all ecological
factors and not by any individual factor. All these factors interact and are interrelated to each
other, forming a complex process. Variations in one may affect the other in measurable
quantities.
48
Figure 3. Simplified patterns of energy and nutrient exchange in the Schirmacher Oasis.
49
impacts than is the land-based ecosystem. The biotic component of the oasis is expressed in
terms of their energy exchanges and trophic position such as producers, consumers and
decomposers.
3.2.1. Producers
The producers of the SO are comprised of all non vascular plants. Based on the study and
collection of all the vegetated sites in the 14 study areas of SO, a total of 34 species were
recorded consisting of mostly lichens, algae and mosses. The Oasis is represented by
individual rare patches of lichens on rocky substrates and by moss mats on silt. A total of
about 19 species of lichen have been reported in the Oasis (Pandey and Upreti 2000). In the
present study 17 species of lichens were recorded. Diatoms were found to be present in waters
of lakes of the oases. The algal-flora of lakes in the oasis consists of about 11 species and
most of them belong to the phytoplanktonic groups dominated by cyanobacteria. On the land,
the producers such as lichen, mosses and algae were found mostly on damp soils, surface of
rocks, and beneath the rock surface. Wherever moisture availability is high the populations of
producers are also high. As the primary productivity is very less, they may support a small
fraction of living consumers. Description of the species found in the SO performing the roles
of primary producers are:
(i) Algae. The algal species at the SO occupied distinct levels of habitats; (a) in
association with mosses (b) on damp soils (c) on quartz rocks in the water pools and
(d) lake bottoms. The submerged rocks, which are directly exposed to sunlight,
favored the growth of thin reddish brown to blue-green encrustation. These crusts
were composed of both N2 and non-N2-fixing species. The cyanobacterial patches
only covered with high amount of mucilage were abundant on the soil surface near
the edge of the stream and were also observed in the depressions created on turning
the small rocks and stones in slow running streams. Glacier does not support growth
of algae and cyanobacteria. The pond and upper stream support less growth of algae
and cyanobacteria. However, their growth was abundant and readily visible on the
surface of the rocks, boulders and weathered soil of the middle portion of the stream
of site no. 2. Algae are also found under rocks, particularly light-coloured quartz
stones, where the microclimate is more favourable than in the surrounding sand or
soil. This strategy enables them to scrape a humble living in this harsh environment.
The species richness of algae was highest in the streams. Analysis of SO lake water
showed the presence of Oscillatoria, Chroococcus, Synechocystis and some diatom
species. Phormidium frigidum constitutes an important and dominant species of algal
flora, which is non-N2-fixing species. Table 4. shows the different types of algae
found at the SO. Majority of the algae recorded at the SO possess tough and collared
mucilage. These properties of algae may help in two possible ways (a) light filtration
and (b) increasing water retention capacity. Komarek and Ruzicka (1966) found that
the ice-covered lakes of SO were highly productive due to algae. The main
component of flora is the Phormidium species, which forms large fan shaped
colonies supersaturating the water with oxygen.
(ii) Lichens. Lichens are the most widespread flora of the SO. Lichens such as
Rhizocarpon flavum, Acarospora gwynnii, Xanthoria elegans, Buellia pallida,
50
3.2.2. Consumers
The real consumers of the oasis are mainly the primary consumers composed of
invertebrate micro-fauna. These micro-fauna have adapted to the extreme living conditions
throughout the years. However, some of the larger birds (Skua, Snow Petrels, Storm Petrels
and stray Penguins) migrate to this region only during the local summer. All the birds depend
on the sea for their food except Skua, which also feeds on the dead remains of Penguins,
Petrel and their chicks. Thus, Skua acts as a detritivore and scavenger. It may be noted that
these birds except Penguins, migrate purposely towards the oasis region, while Penguins
come by mistake, losing their sense of direction. Instead of moving towards the coast in
search of food, they move towards the continent. The death of such Penguins serves the
purpose of the scavenger bird Skua and the remains add to nutrient pool of the oasis. The
invertebrate terrestrial fauna of SO mainly inhabited in the soil and in vegetation. They range
from protozoa (single-celled creatures), rotifers, tardigrades and nematodes to arthropods
51
(mainly mites and springtails, midge). The largest invertebrate found is the wingless midge
(Belgica antarctica), which grows to 12 mm long. The distribution of tardigrades, nematodes
and rotifers are more near surface soil area. The occurrence of micro fauna on the top layers
of the soil may be due to a slight rise in temperature due to sunshine, availability of food etc.
They may also migrate up and down in the soil.
Further studies might reveal the local variations due to temperature and light difference
the occurrence of these fauna in the soil. Heywood (1979) has also shown that many
invertebrates are plant feeders. It may be stated here, that the species reported in 8 lakes and 4
swampy areas of SO are those that are distributed widely in Antarctic lakes with similar
behavioural pattern that is thriving well in sediments rich in micro flora and organic matter.
Densities of these micro fauna vary from 20,000 to more than 14 million animals per m2
(Bonner and Walton, 1985). Ingole et al., (1987) observed a maximum tardigrade density of
140 per m2 and 272 per m2 of nematodes at Zub Lake.
However, a maximum of 35 per m2 of Tardigrades, 21 per m2 of nematodes and 10 per
2
m of rotifers were observed in the present study, which is much lower than at the bottom of
the lake in SO. The low density of micro-fauna in the present study may be due to less food
availability (moss growth) on the land. Population density of these micro-fauna is reported to
show variations with light, temperature, level of blizzard, relative humidity and food, these
being the major determinants (Fleeger and Hummon, 1975; Morgan, 1977). All of the
Antarctic animals have adapted to life in extremely cold conditions. The springtails and mites
live under rocks in the SO. A brief description of the primary consumers of SO is provided as
follows.
Table 4. Taxonomic assessment, identification, habitats descriptions and the dispersal
potential of the algal communities of Schirmacher Oasis, East Antarctica
S No.
Habitat
descriptions
Shapes and
sizes (m),
color
Forms of
possible
dispersal
Comments
Class:
Cyanophyceae
Order:
Nostocales
Calothrix gracilis
Lyngbya
aeustuarii
Nostoc commune
Oscillatoria
limosa
Phormidium
frigidum
Schizithrix
Order:
Chroococcales
Aphanocapsa
Aphanothece
nidulans
Gloeocapsa
kuetzingiana
Found in all
the sites; fresh
water lakes,
streams,
swamps,
ponds etc.
association
with lichen
and mosses;
forming green
scum; black
epilithic crusts
on rock
surfaces
Ellipsoidal,
Spherical,
irregular;
motile or non
motile; 1-90;
both
unicellular or
multicellular;
colonial
Independent
dispersal of
spores and
cysts may
occur;
vegetative
cells, spores
and cysts
Generally
aggregated,
chlorophyll
present, thallus
undifferentiated
thalloid plant body
52
Habitat
descriptions
Shapes and
sizes (m),
color
Forms of
possible
dispersal
Comments
Spherical,
ellipsoidal,
irregular
1-80
Same
Same
Order:
Chlorococcales
Chlorococcum
3
Class:
Baccilariophycea
e
Order:
Pennales
Hantzschia
Pinnularia
borealis
Lichen class,
order and species
Sites, Habitat
descriptions and date
Shapes and
sizes (m),
colour
Spherical,
irregular;
35-70;
yellowish
green
Forms of
possible
dispersal
Independent
dispersal of
ascospores;
Soredia, Isidia
Comments
Acarospora
A. gwynnii
A. williamsi
Alectoria
A.minuscule
Very common on
shady place, moraine
and rocks; generally
absent in habitats
close to melt water;
Near Russian and
Indian Stations
Cracks and surface
of rocks, rocks inside
water
Same
Same
Moraine, amongst
pebbles, rocks
Same
Same
Carbonea
C. capsulata
Grow on rocks of
small depressions,
Near Russian station
along the lake.
White mat
with
compressed
areolate
Same
Lecidea
L.cancariformis
Same;
black
apothecia
Same
Lecanora
L.fuscobrumnea
Very common,
endolithic lichen,
grow in dry rocky
surface, mostly on
the leeward sides of
rocks, along the dried
stream
Grow on rocks,
sheltered place,
Same; Fruticose;
forms varicose
masses;
filamentous
Same; umbilicate
thallus; crustose
and saxicolous
Same;
amorphous
thallus; apothecia
on the tops and
sides of the
stipes
Same
Buellia
B.pallida
yellowdark brown
disc,
Same
Squamulose
areolate and
crustose; thallus
fragments very
common
Same; lichen
with crowded
mats of
apothecia
Lichen class,
order and species
Sites, Habitat
descriptions and date
Lepraria
Lepraria
membranacea
Physcia
P.caesia
Polycauliona
P. murrayi
Porpidia
P.species
Rhizocarpon
R.flavum
R.species
10
Rinodina
R.Species
11
Umbilicaria
U.aprina
U.decussata
12
Xanthoria
X.elegans
Stones, pebbles,
Rocks; leeward sides
of rocks
Rocks, stones,
pebbles; streams,
ponds, lakes
53
Shapes and
sizes (m),
colour
Powdery,
Yellow
colored
Forms of
possible
dispersal
Same
Comments
Same;
whitish,
Same
Same; foliose
lichen
Orangeblack
circular
hapteron
Same;
black
apothecia
Same;
yellowblack and
white-black
colored
Same;
black
granular
Same
Same; fruticose
lichen; erect tufts
Same
Same
Same
Same
Same
Same; varicose
lichen
Thallus 0.1
cm to 18
cm
diameter
Same
Same; foliose
lichen;
cosmopolitan
Same; red
lichen
Same
Same; lobate
lichen
Same
Mosses family,
Genus and species
Sites, Habitat
descriptions and date
1.
Bryaceae
Bryum
argenteum(Hedw.)
Bryum
pseudotriquetreum
(Hedw.)
Globose,
tetrahedral,
Variable in
length;
10-25; yellowish
brown to
brownish, red
stem, deep green
Forms of
possible
dispersal
Spores,
Gemmae
Comments
Sporophyte
present;
seta 1.5-4.5
cm long;
whitish
colour;
capsulated;
vegetative
fragments
dispersal
observed
54
S
No.
Mosses family,
Genus and species
Sites, Habitat
descriptions and date
2.
Pottiaceae
Bryoerythrophyllu
m recurviroste
(Hedw.)
Pottia cf. heimii
(Hedw)
3.
Ditrichaceae
Cedratondon
purpureus (Hedw.)
Common; Same as Sl
No. 1
4.
Grimmiaceae
Grimmia sps.
Common; Same as S
No. 1.
Globose and
tetrahedral;
10-25; reddish
green to brown,
yellowish green
to green, reddish
nerves
Globose,
tetrahedral,
Variable in
length;
10-25; brownish
green, yellow,
reddish
Globose,
tetrahedral,
Variable in
length;
6-20; brownish
green, green
turfs
Forms of
possible
dispersal
Gemmae,
vegetative
fragments
Comments
Gemmae,
vegetative
fragments
Non
Sporophyte
vegetative
fragments
dispersal
observed
Gemmae,
vegetative
fragments
Non
Sporophyte;
vegetative
fragments
dispersal
observed
NonSporophyte;
vegetative
fragments
dispersal
observed.
55
and moss dwelling forms. However, earlier workers found that several species are
endemic to SO.
(ii) Rotifer. The Rotifers are a group of small usually microscopic, Pseudocoelomate
animals. They are also called Rotatoria or wheel animalcules. Only one species
viz., Philidina gregaria has been observed and collected from SO.
(iii) Tardigrada. Tardigrades have well defined head and four trunk segments with 4
pairs of short legs bearing claws, which are used to walk along underwater surfaces.
Tardigrades have the ability to hibernate (cryptobiosis) in which they can survive
extreme thermal condition, exposure to highly toxic chemicals, drying out, etc. In
this process, they are capable of withstanding very cold condition (-900 C) by passing
into a state of very low metabolic activity. This phenomenon is also referred to as
anabiosis. The body surface is covered with ornamented plates, sometimes bearing
spines or hairs. Many species have eyes. The mouth is terminal or ventroterminal.
Sexes are separate and the females are oviparous. Development is direct, the cuticle
being moulted. Two species of tardigrades are found in SO, namely Hypsibius
ckhilenesis and Macrobiotus polaris. The ability to tolerate severe desiccation,
anhydrobiosis survival etc are the advantages for the widespread distribution of many
species of tardigrades. In spite of the short summer season tardigrades multiply
quickly and become very abundant. They reproduce by parthenogenesis.
(iv) Nematoda. Three species of nematodes are found in SO viz. Tylenchorhychus sp.,
Dorylaimoides sp and Rhabditis. Hazra (1994) recorded 5 genera from SO for the
first time. The genus Tylenchorhynchus and Dorylaimoides, which occur widespread
in the Indian continent, might have been transported along with agricultural products
to the SO especially in the Zub lake area (Venkataraman 1998).
(v) Springtail (Collembola). Two species out of two families viz., Isotomidae and
Entomobryidae were observed from SO. Isotomidae is larger and have slender body,
they can jump actively, whereas smaller type Entomobryidae is dorso-ventrally
flattened with broad oval abdomen and moves slow to hide in the soil. The body
segments are more obvious in larger type. The springtails were collected from soil
with micro-plants. They mainly inhabited the soils where little growth of plants is
possible. The population densities of larger and smaller types of springtails were
counted as 6 to 28 and 12 to 56, respectively in 100 gm of soil sample. So far 20
species of springtails have been reported from the Antarctic continent (Laws, 1977).
(vi) Mites. Mites, which belong to the spider family, are the commonest land animals. It
is the worlds most southerly indigenous animal found as far south as 850 S. Two
species of mites viz., Tyrophagus sp and another unidentified species of family
Scutacaridae from SO have been reported by earlier workers (Mitra, 1999). Many of
the mites avoid freezing by a physical process known as super-cooling, whereby
their body fluids are maintained in a liquid state in temperatures below their normal
freezing point (Pan and Shimada 1991). Their ability to synthesize glycerol,
antifreeze, enables them to survive temperatures of 350 C (Block et al., 1994). The
population varied from 7 to 76 in 100 gm soil. Acarina represents the principal
Arthropod group and these mites are large and relatively well-known group in the
region.
(vii)
Insects. Compared to other regions, insects are scarce and much smaller in
size over the oasis. The type of living habitat for most of them has been found to be
56
Microconsumers
(species)
Habitat descriptions
Protozoa
Rotifer
Tardigrada
Nematoda
Springtail
(Collembola)
Insects
Mites
Non-planktonic; Found in
aquatic mosses and algae,
mud, ponds and lakes.
Active tardigrades found
even in droplets and film
of water on terrestrial wet
mosses.
Not common; found in
soils, plants and dead
organic materials,
terrestrial moist soil,
moss-fresh-water etc.
Found mostly in soil;
damped place with small
amount of micro flora
Scarce; Parasitic; found
mainly inside the
stations; birds feathers
etc.
Commonest land animals
Comments
Small microscopic,
mm long;
reddish color.
Also known as
rotatoria or
wheel
animalcules
Commonly
known as water
bear;
Cryptobiosis;
reproduce by
parthenogenesis
About 1 mm long
About 0.6 mm
long, 0.03 mm
wide and weighs
about 0.55
micrograms
About 0.65mm to
1.25 mm long;
white in color
Smaller compare to
mainland parasites
About 0.3 mm
long; whitish in
color; oval shape
with dorsally
convex body
Worlds most
southernly
indigenous
animal; process
of "supercooling"
57
58
3.2.3. Decomposers
At the present study the decomposer assessment could not be undertaken. However, some
earlier workers have reported the following microorganisms. Bacteria, yeasts and fungi: The
dominant species of bacteria present in SO are the gram-positive rod identified as
Pseudomonas fluoresceus, P. putida and P. syringae and gram-positive cocci as Micrococcus
genus Arthrobacter (Shivaji et al., 1989a, 1989b). Matondkar and Prabhu (1986) reported
bacterial counts in peripherial lake sediments in January 1985 to range from 21x105 numbers
g-1 of dry sediment. In loam and moss soil, their concentration varied from 12x102 to 2.5 x
103 and 10 x 102 to 1.2 x 103 per g dry weight respectively. Moss algae and moist soil were
rich in bacterial counts as compared to dry sand. Evans et al., 1997 has reported bacterial
counts to vary from (1 to 25) 106 per g dry weight in some of the Antarctic lake sediments
and majority of the bacteria as psychrotrophs. On the other hand phylogeny of obligately
anaerobic, coiled bacterium from Ace Lake, Antarctica, was isolated by Franzmann and
Rohde, (1991); Franzmann and Dobson (1993), which they believed to have been introduced
by humans. Franzmann et al., (1991) also found that Carnobacterium funditum and
Carnobacterium alterfunditum are psychrotrophic, lactic acid-producing bacteria found at
anoxic waters in Ace Lake, Antarctica. Matondkar and Prabhu (1986) studied the effect of
temperature on bacterial population of SO, and found that temperature has minimal effect on
these microorganisms indicating their high degree of tolerance to low temperature. Shivaji, et
al., 1989c isolated fungal populations and found Penicilium species, consisting of P.
olivicolor, P. corylophulum, P. viridicatum, P. chrysogenum, P. waksmanii, and P.
camemberti to dominate the fungi. Few colonies of Fusarium oxysporum and Paecilumyces
variotii were also reported. Similarly, in the soil samples of SO, 8 strains of yeasts
Rhodotorula rubra, one Bullera alba, one diamorphic Candida humicola and one C. famata
and the remaining two tentatively identified as C. ingeniosa and C. auricularia was reported
(Ray et al., 1989). In the dry valleys elsewhere in Antarctica various yeast strains such as
Cryptococcus, Candida, Rhodotorula and others have been reported (Vishniac and Kurtzman,
1992; Vishniac 1993).
59
The carbohydrates produced by the algae and mosses through photosynthesis are the
main source of food for the primary consumers like mites, tardigrades, protozoans, rotifers
and midges etc.
More information is needed on the actual decomposition process in the Antarctic
ecosystem where temperature is very low. The primary production is very little and as such
the proportion of plant biomass passing into detritus component is presumably quite low as
most of the primary production is grazed by the herbivores. Detailed investigations are
required to understand the energy flow and nutrient cycling through the terrestrial ecosystem
at SO through the simple food chain. During the Antarctic winter when it is dark, there is no
photosynthesis by many of the producers. The plant eating primary consumers living on algae
grew only during the summer.
60
Vegetation
Groups
Name of Species
Frequency
(%)
Algae
II
Lichen
III
Mosses
Lyngbya aeustuarii
Nostoc commune
Oscillatoria limosa
Phormidium fragile
Aphanothece nidulans
Chlorococcum
Pinnularia borealis
Acarospora gwynnii
Buellia pallida
Carbonea capsulata
Lecidea cancariformis
Lepraria membranacea
Physcia caesia
Polycauliona murrayi
Rhizocarpon flavum
Rinodina Species
Umbilicaria aprina
Umbilicaria decussata
X.elegans
Bryum argenteum
Bryum pseuotriquetreum
Bryoerythrophyllum
Recurviroste
Cedratondon purpureus
Grimmia sps.
Unknown
60
60
80
80
20
40
40
40
40
20
30
20
10
30
20
10
30
30
10
60
20
10
Frequency class
To which species
belongs
C
C
D
D
A
B
B
B
B
A
B
A
A
B
A
A
B
B
A
C
A
A
20
20
70
A
A
D
IV
Unknown
The above-calculated values are plotted and compared with the normal frequency classes
(Figure 5.a and b). From the figure 5.b. it is clear that values of frequency classes B, C and D
of figure 5.b. is comparatively higher than the respective values in normal frequency diagram
(Figure 5.a). Thus, the plant community of the SO is heterogeneous in character. It can also
be mentioned here that the value of frequency class E in figure 5.b is nil. Thus, none of the
61
species showed 80-100% constancy. This indicates that the SO environment does not provide
suitable habitat for any species to occur homogenously. The heterogeneity is linked to
microenvironments in the oasis providing congenial conditions for growth, both spatially and
temporally. Also from table 8 it is found that the dominant algal species are Oscillatoria
limosa and Phormidium fragile both of which are blue green algae, the moss species are
dominated by Bryum argenteum and the lichen species by Acarospora gwynnii. It is
important to note here that amongst all the plant groups, algal species dominate, whereas, the
occurrences of moss and lichen species is lesser, which could be because the study sites are
mostly aquatic in nature and it is true that out of 14 selected sampling 8 sites are just the
banks of lakes, two dried lakes and four swampy areas. Some unknown species also occurred,
which seemed to be some moss in close association with some lichen or algae, and could not
be identified. It is very interesting to note that all the algal mats were dominantly consisting
of cyanobacteria or blue green algae. Heterocystous cyanobacteria capable of independent
nitrogen fixation like Nostoc Commune were mostly present on ice surface and lake bank over
snowmelt. Pinnate diatom Pinnularia was found on surface layers of lift-off and often seen
sliding between blue green algal mats. The only green algae found in the present transect
study was Chlorococcum. This is in contrast to many other reports from snow regions where
Chlorophyta members are present in abundance. Earlier Ling and Seppelt (1993) reported
Chloromonas Subroleosa belonging to chlorophyta from Antarctica, which gave a red color
to the snow attributed to the development of pigment in the species.
Figure 5. A and B. Shows the Raunkiaers normal frequency diagram and the frequency diagram of the
Schirmacher Oasis community. Frequency classes B, C and D of diagram B are relatively higher than
A, thus the community is heterogeneous.
62
Rare occurrence of green algae and dominance of blue green algae in the SO of
Antarctica suggests that these cyanobacterial species are better adapted to the prevailing
extremities of climatic conditions in Antarctica. The prokaryotic cell wall struture consist of
diaminopimellic acid and muramic acid presumably provides better resistance to the cells. It
also seems likely that the green algae are not so efficient in withstanding the freezing and
thawing. In an experiment conducted by Bidigare et al., (1993), samples of red and green
Chlamydomonas (Chlorophyta) when exposed to freezing and thawing resulted in lysis of the
green cells whereas the red cells survived, which was due to increased membrane fluidity.
Cyanobacteria containing phyco-cyanin (blue pigment) and phyco-erythrin (red pigment)
seem to have more resistance to the stress conditions of Antarctica.
Site No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
Biomass (gm-2)
Fresh weight
11.5
15.35
12.5
9.54
14.5
26.7
8.5
21.75
7.5
21.75
17.45
8.5
11.6
21.8
Dry Weight
2.8
3.25
2.95
2.2
3.45
7.25
1
6.5
1.2
6.2
4.65
1.2
2.46
5.3
Moisture Content %
310.71
372.31
323.73
333.64
320.29
268.28
750
234.62
525
250.81
275.27
608.33
371.54
311.32
17.5
20.31
18.44
13.75
21.56
45.31
6.25
40.63
7.5
38.75
29.06
7.5
15.38
33.13
Mean (1 S.E)
14.92 ( 4.30)
3.60 ( 4.32)
375.42 ( 4.31)
22.50 ( 4.32)
63
It must, however, be mentioned here, that these values of NPP are highly biased because
they correspond to the study sites where conditions are favourable for growth. Even during
summer, there are large areas bearing no vegetation at all. The present estimates of NPP
therefore give an estimate of the potential of production of SO under the prevailing
conditions. The average NPP value of the SO (22.5 gm-2), when compared to other stressful
ecosystems of the world is still found to be very little.
Table 10. Physicochemical analyses of the various water, soils and biological samples and the atmospheric conditions
of the particular dates
S.
No
1
2
3
4
Ecological
Parameters
Surface Air
Temp. (0C)
MSL
Pressure
(mb)
Wind Speed
(m/s)
UV-B
(MED/Hr)
IX
XII
Average
VI
-2.63
-2.63
-2.63
0.2
0.2
0.2
2.5
2.5
2.5
-2.0
-2.0
-2.0
-3.2
-3.2
-0.87
986.57
986.57
986.57
978.96
978.96
978.96
981.25
981.25
981.25
981.9
981.9
981.9
986.3
986.3
982.76
3.77
3.77
3.77
5.76
5.76
5.76
4.50
4.50
4.50
4.11
4.11
4.11
3.08
3.08
4.32
0.798
0.798
0.798
2.282
2.282
2.282
2.1
2.1
2.1
0.946
0.946
0.946
1.718
1.718
1.55
DO mg l-1
8.2
7.8
7.9
6.1
7.6
6.6
8.4
8.1
8.2
6.5
7.9
6.2
7.8
7.6
7.5
6
7
8
9
10
pH
PO4 mg l-1
NH4 mg l-1
NO2 mg l-1
NO3 mg l-1
7.2
0.16
0.36
0.45
0.56
7.8
0.42
0.01
0.06
0.78
7.4
0.24
0.24
0.22
0.98
8.2
0.12
0.12
0.18
7.3
0.04
0.12
0.32
0.12
7.6
0.56
0.03
0.14
0.32
7.5
0.32
0.34
0.46
0.51
6.9
1.28
0.93
0.94
0.94
7.7
0.61
0.56
0.76
0.12
7.8
0.93
0.01
0.07
0.23
7.8
0.07
0.23
0.82
2.40
7.1
1.26
0.99
0.19
2.20
7.0
0.07
0.17
0.21
0.09
7.4
0.08
0.06
0.05
0.16
7.5
0.44
0.31
0.3
0.69
11
Ca mg l-1
22.6
2.1
3.1
25.0
32.0
14.3
10.5
37.5
29.0
2.5
6.4
48.5
10.2
2.2
17.6
12
Mg mg l-1
Chlorides
mg l-1
Chlorophyll
mg m-3
Primary
Productivity
mgC m-3 hr-
4.2
3.1
1.3
3.0
16.4
2.5
3.5
15.2
2.3
2.4
1.1
9.3
1.3
1.1
4.8
0.016
0.022
0.018
0.022
0.011
0.008
0.041
0.052
0.021
0.026
0.056
0.052
0.001
0.000
0.0
0.264
0.632
0.046
0.050
0.457
0.132
0.241
0.060
0.415
0.122
0.711
0.910
0.058
0.260
0.31
0.72
4.15
1.13
0.62
3.26
2.70
1.13
4.26
1.85
2.80
0.65
4.33
0.89
0.55
2.074
1.36
1.77
1.43
1.36
1.70
1.63
1.43
1.77
1.57
1.63
1.36
2.58
1.36
1.16
1.582
13
14
15
16
Organic
Carbon %
S.
No
17
18
Ecological
Parameters
Sediment
Organism
density
individuals
m-2
Dehydrogen
ase activity
g/ g soil/24
hr
IX
XII
Average
VI
1730
520
2205
1100
1115
1680
1750
1820
1050
1250
410
430
1630
380
1219
0.007
0.011
0.008
0.003
0.01
0.009
0.009
0.012
0.011
0.009
0.003
0.016
0.008
0.001
0.008
66
67
that of marginal soils in sub-tropics (Malik et al., 1995). The DHA value shows a relationship
with soil organic carbon. The low DHA in SO soils indicates that there is very limited
microbial activity in the soil, which in turn, affects the decomposition process and Cmineralization. Relatively higher DHA values on the site 12 may be attributed to the inputs of
the organics matter coming from the Russian Station Novolazarevskaya (Gajananda 2007).
3.6.1. Temperature
Temperature plays a key role in the survival, growth, locomotory activity, life cycle,
oxygen consumption rates, water balance, osmoregulatory ability, psychrophilic forms, fatty
acid synthesis and other metabolic characters of plants and animal (Block et al 1994; Klok
and Chown 1997; Moller and Dreyfuss 1996).
68
A perusal of the literatures shows a wide range of the effects of temperature on the flora
and fauna of Antarctica. Many earlier workers had studied various aspects of temperature and
their roles in determining the organisms survival, evolution of super cooling ability and
habitat distribution etc.
Booth (1990) used various climatological data to describe the climatic requirements of
plants and for numerical methods and models to indicate the decisive climatic factors for
plant distribution. In another study of temperature dependences of three species of free-living
Antarctic fellfield nematodes exhibited differing degrees of both strategies of cold-hardiness,
freeze-tolerance and freeze-avoidance (Pickup 1990). The adaptation and acclimation of
growth and photosynthesis of some Antarctic freshwater algae to low temperatures had been
studied (Nagashima et al 1993; Eggert and Wiencke 2000).
Both low temperature and water are limiting factors in the Antarctic terrestrial
environment; these two factors have profound effect on the rate of photosynthetic response to
the Antarctic moss Polytrichum alpestre to low temperatures (Kennedy 1993a-b). Franzmann
(1997) found that cells of Methanogenium frigidum isolated from the perennially cold, anoxic
hypolimnion of Ace Lake in the Vestfold Hills were psychrophilic, which exhibit most rapid
growth at 150 C and no growth at temperatures above 18 to 200 C. The growth and
consumption rates of bacteriovorous Antarctic naked marine amoebae grew at temperatures
down to 20 C and showed optimal growth at 20 C and the growth efficiencies were low,
typically <2%, which suggest that amoebae incur high energetic costs for life at low
temperatures (Mayes et al 1997). Crustose lichen Buellia frigida growing on rock at Cape
Geology were well adapted to the combination of hydration, low temperatures and strong
light (Kappen et al 1998). Studies carried out on the cyanobacterium Nostoc commune by
Kashyap et al (1991) over SO shows the influences of temperature on the nitrogen fixation.
Matondkar and Prabhu (1986) also found that in SO the effect of temperature on bacterial
populations is significant. At SO, the animals recorded were all of very small size. The small
size also seems to be an adaptation to the prevailing low temperature. Todd and Block (1997)
and Pickup (1990) showed greater super-cooling ability in small-sized classes of insects due
to accumulation of cryoprotectants.
Freezing intolerance is much more widespread throughout the invertebrates than freezing
tolerance, but that switching from being tolerant to intolerant of body ice formation may
occur in natural populations under snow cover (Block et al., 1994). In Antarctica the
organisms are eurythermal, which can survive temperature in a very wide range. One good
example was given by Nicolaus et al (1998), who found that bacterium Alicyclobacilli
isolated from an unexplored geothermal soil in Antarctica: Mount Rittmann, were
thermoacidophilic, showing an optimum temperature of -630 C and an optimum pH of 3.5 to
4.0. These bacteria were able to grow in the temperature range -45 to 700 C. Thus, in
Antarctica both cold tolerant and thermophilic organisms can survive. Thus, from the above
discussions it is clear that temperature plays a great role in all the life forms of plants and
animals.
It seems to affect shape, size, and volume, short or long life cycles etc. The plants as well
as animals present at SO have adaptations to face the low temperature and are tolerant to
freezing. Also, the snow present at Antarctica plays great role in providing suitable niches to
the organisms through its various physical properties. The data of Maitri and the
Novolazarevskaya are complimentary to each other giving an opportunity to study long-term
variation or time series analysis of both temperature and pressure.
69
Figure 6. Seasonal average temperature from 1961 to 2001 recorded at Novolazarevskaya, Antarctica.
Full data sets of annual temperature (dark squares) with summer values highlighted as open squares.
Solid trend lines represent annual and dashed lines are of summer only.
Figure 6 shows the temperature trend of the Novolazarevskaya station for the last four
decades. The data pertains to the year 1961 to 2001. The monthly average surface air
temperature observed at this station shows that the temperature has significantly increased
during the last four decades by [1.11 0C (P<0.01)], which is at the rate of about 0.27 0C per
decade. The summer (December-February) of this period also shows a significant rise in
temperature by about [0.99 0C (P=0.05)], which is at the rate of about 0.24 0C per decade
increase. In the year February 1996 a widespread warming over east Antarctica was reported
(Gajananda et al., 2002d). Hence, in the SO, the surface air temperature enhancement will
certainly affect the function of ecosystem dynamics in future.
70
Figure 7. Monthly average msl pressure observed at Novolazarevskaya from 1961 to 2001.
In Antarctica specially, pressure plays a vital role in the development and maintenance of
flora and fauna indirectly by affecting cloud formation, precipitation etc. The high and lowpressure systems around the circumpolar trough develop in severe cyclone. These cyclones
push towards the continent and result in the formation of clouds, moisture, warmer air and
precipitation over the continent periphery (Bromwich 1997). The cyclones also churn the
ocean, thereby mixing the water with atmospheric oxygen, resulting into very high production
of Southern ocean. Likewise, over the continent periphery, warmer air, precipitation, churning
of lake water etc. gives a better habitat for plants and animals.
71
Figure 8. Albedo (%) of various vegetation types and other surfaces values.
72
The amount of solar radiation reflected by the soil surface or by vegetation varies
considerably. Green plant cover reflects little of the visual spectrum but much above 700 nm
(near infra-red). Albedo (%) of various vegetation types and other surfaces values are given in
figure 8. (after Stoutjesdijk and Barkman, 1991). From this figure, we see that water has a low
albedo, but at low solar elevations, specular reflection strongly increases the albedo. The
albedo of fresh snow is the highest found on earth and approached the albedo of an ideal
white surface. The net radiation is the amount of energy available for the warming of soil and
air and for evaporation. Clearly, the net radiation will differ markedly from point to point; a
bare surface which is greatly warmed by the sun will receive a low net radiation and,
conversely, a strongly transpiring crop with the same albedo as the bare surface, remaining
cool, will have a high net radiation. Also, when there is a strong wind, the net radiation
received by bare sand will be greater than under still conditions, when the surface is
considerably warmed up. The warming of the atmosphere by the soil surface can be extended
to hundreds of meters in the course of a sunny day. Naturally, thermal convection is
dependent on the intensity of solar radiation (at night there is no convection at all) and on the
characteristics of the landscape: a strong convection above dry ground, but less above
vegetation using a large part of the energy received for evaporation.
73
Figure 9. Graph showing the maximum daily values during the noontime for the period 1st February,
1998 to 31st January, 2000 observed over Maitri, Antarctica.
A comprehensive long term program involving the study of ecosystem under both
prolonged low UV-B conditions and high UV-B conditions should be drawn, so as to
understand the relationship and scientific mechanisms through which the ecosystem of SO
has survived.
Although many studies, but not all (Tosserams and Rozema 1995, Gehrke et al. 1995),
suggest that photosynthesis and plant growth can be reduced by elevated levels of UV-B
radiation, data on plant biomass and rates of net primary productivity, i.e. the standing stock
of living matter expressed as g dry weight per unit ground area, and growth expressed as g
dry weight per unit ground area per unit time, are difficult to find for natural ecosystems and
forests. However, a review (Rozema et al. 1997a) suggests that there is little impact of UV-B
radiation on the primary production of terrestrial ecosystems.
The high doses of UV-B radiations in the Antarctic ecosystem also result in strong photooxidation of chlorophyll-a of green algae causing damage to the algal cells whereas other
pigments are more protective against UV. This could be another reason why the species
dominating the SO ecosystem consisted of cyanobacteria. Lichens, which are the symbiotic
associations of cyanobacteria and fungi mainly, occupied hilly and rocky terrians. Due to
their highly adaptive features of nutrient absorption by heterotrophic fungi and photosynthesis
by algal partner, they are the common pioneer species reported for starting autotrophic
succession on bare rocks.
3.6.7. Snow
Snow cover changes the thermal budget at the surface of the earth. Fresh snow reflects
solar radiation, not only the visible light but also shorter and longer wave radiation, up to
95% (Figure 8). Older snow has a lower reflection because of the snow turning into ice and
due to accumulation of dust. However, for long-wave (heat) radiation snow acts as an ideal
black surface, with absorption of 99.5 % and only 0.5% reflected. Older snow partly turns
into ice and becomes more transparent. A fresh snow cover of 20 cm transmits only 3% of the
solar radiation between 400 and 700 nm, but an older snow cover of the same depth transmits
74
13% (Geiger 1965). Under icy snow a sort of greenhouse effect may occur. In Antarctica
temperatures of +130 C were measured on lichens growing under an icy snow cover, with an
air temperature of 90 C (Lange and Kappen 1972).
Because of the small amount of heat from the soil during a clear and still night snow can
cool off rapidly at the surface, causing very low night temperatures. Moreover, high radiation
intensities during the day (both from direct and reflected sunlight) may lead to relatively high
temperatures in the middle of the day. As a result, plants growing above the snow may be
subjected to very large temperature differences, with a damaging effect.
A particular characteristic of snow is its insulating effect, protecting soil and vegetation
from frost. On top of the snow cover it is much colder than on the bare ground; underneath
the snow it is much warmer. Many plants in alpine and arctic/Antarctic environments with
winter temperatures of 20 to 50 0 C, would be killed in temperate regions because of the
severe frost with no protecting snow cover. Under a long-lasting snow covers many animals;
both vertebrates and invertebrates can remain active for most of the winter (Merriam and
Caldwell 1983).
Lichens are photosynthetically active when they receive a great deal of light reflected by
the snow on the ground. Creveld (1981) found that during winter lichen receives the highest
light intensities because of snow reflection and called this adaptation as cheimophotophytic
(cheimoon = winter). This is also true for Antarctic lichens.
Snow offers an effective protection against low temperatures; on the other hand little
solar radiation is passing through a thick layer of snow. Another ecological aspect of snow is
that the snowmelts on the soil may be saturated with water. Thickness of the snow cover and
time of melting are often varied spatially according to fixed patterns, which can be correlated
with vegetation patterns found in the summer (Friedel 1961).
Sheltered depressions and spaces behind obstacles receive a great deal of snow. In the SO
snow remains longer on north slopes than on south ones. Variation in snow cover ranges from
very short duration with strong frost and desiccating winter storms (Blizzards), to a relatively
long duration, especially in SO. Stoutjesdijk and Barkman (1991) found that if the soil is
snow-free for less than three weeks, no vegetation will develop. If the snow-free period lasts
at least three weeks the moss dominated. If the snow-free period is still longer, and the winter
is cold, the limiting effect of frost on the vegetation is considerable.
Clearly, lichens growing in snow-free places in high mountains and polar areas have to
tolerate very low temperatures. In the SO the average summer temperature is 9.83 0C, the
average winter temperature is 300C. Lichens occurring here tolerate temperatures of 750C,
even in wet conditions (according to laboratory experiments of some workers). They can
assimilate at 20 0C and have their photosynthetic optimum at low light intensities and very
low temperatures of 0-50 C (Lange and Kappen 1972; Gjessing and Ovstedal 1989). The dark
colour of lichens above the snow cover may be related to their heat requirement.
Solar radiation, because of the high reflection from snow, has little influence on
snowmelt. As soon as objects appear above the snow, plants or stones, melting proceeds
much faster. These objects release heat through conduction and radiate heat, which is fully
absorbed by the snow. In this way holes arise in the snow around patches of mosses, algaes
and lichens, providing the essential water for these plants.
The extreme low temperature of Antarctica resulting in the thick snow cover forms an
important part of the climate system and affects greatly the ecology of the region. The snow
cover seems to mediate the SO ecosystem in the following ways:
75
a) The snow has a great potential of acting as an energy bank by storing latent heat of
fusion and sublimation and crystal bonding forces (Gubler, 1985). The intake and
release of energy throughout the year seems to be responsible for the variability in
habitat conditions leading to heterogeneity of the ecosystem composition at the SO.
b) The snow has a porous medium with large air content with a high insulation capacity.
The insulation plays an important role in giving protection to the microorganisms.
c) The snow acts as a reservoir of water during the summer and provides habitat and
food resources to the various primitive plant forms, invertebrates and microbes.
Permafrost: Permanently frozen ground underlies some 25% of the earth's land surface,
which is widespread in Antarctica. Permafrost is believed to form gradually over very long
time intervals (105 years), occurring continuously where mean annual air temperature is <-70
C. Along the margins, where it is thin and discontinuous, permafrost degradation can be
monitored. Changes in the permafrost and the overlying summer active layer can have
important effects on hydrological regimes, ecological and geomorphological conditions, and
structures (Koster and Nieuwenhuijzen, 1992; Nelson et al., 1993).
3.6.8. Energy Balance and Water Relations, Dew Formation, Fog and Hoarfrost
During the day high temperatures of the lower air layers promote energy exchange, while
at night the situation is reversed: the colder and heavier air is not easily mixed with higher
warmer layers of air and consequently the atmosphere is stable.
Nyberg (1938) found that at 1 mm above the snow on still clear nights there was an
unmistakable effect of turbulence. As stated above, the vapour pressure profile indicates the
beginning of dew formation, which has been left out of consideration in the energy balance
because it is negligible in energy terms. The occurrence of extremely low minima above bad
conduction soil has received more attention than the absence of such minima above rock
surfaces, although both phenomena are biologically important.
For many organisms, which cannot take up water from the soil, the presence of liquid
water can be very important, from dew or from precipitation. For mosses and lichens dew
may be essential. These organisms can absorb water up to saturation from a dry starting
position and retain this water. In mosses dew is also important for fertilization, allowing the
spermatozoids to swim to the archegonia. Rocks are a similarly extreme environment for
mosses, algae and lichens, as deserts are for higher plants. Some epiphytic species are only
found at the foot of hills and do not grow higher up the glacier where the nocturnal radiation
fog reaches. Dew is of special importance for the micro flora on the surface of leaves of
vascular plants (Ruinen 1961). For many animals, dew is an important source of water.
The relationship between fog and lichen flora in the Antarctica is one of the good
examples. Fog is important as a source of water for organisms, which cannot take up water
from the substrate, such as lichens on rocks. It is also important for lichens to be able to
photosynthesize continuously in the daytime, which requires sufficient light and moist (but
not wet) conditions. Fog here is more favourable to lichens than rain, i.e. especially fog in the
daytime. In climates with mainly night fog and drought during the day lichens do not
thrive, because respiration at night continues and is not compensated for sufficiently by
photosynthesis (a so-called negative photosynthetic balance). In the SO, for example,
the number of days with fog is generally low throughout the years. In the same direction
the epiphytic lichen flora is considerably impoverished. Thus, maps of fog frequency are
76
important if one wants to know whether the fog habitually occurs at night or in the daytime
and affects lichen flora. A prolonged presence of elevated layer in the planetary boundary
layer over SO will enhance fog duration (Gajananda et al., 2002e). Figure 10. shows the
Sodar echogram of the foggy boundary layer in SO during January 1996.
Figure 10. Facsimile pictures of foggy and non-foggy days during 810 January 1996 at Maitri.
Formation of fog started at around 21 : 15 h (UTC).
77
For many organisms, which cannot take up water from the soil, the presence of liquid
water can be very important, from dew or from precipitation. For mosses and lichens fog may
be essential. These organisms can absorb water up to saturation from a dry starting position
and retain this water. In mosses, fog is also important for fertilization, allowing the
spermatozoids to swim to the archegonia.
Clouds form when air is supersaturated with respect to water or ice (Gajananda et al.,
2007). Fog is a potentially significant source of water in the desert environment, and also a far
more predictable source of moisture than rainfall (Gajananda et al., 2007). There is, however,
no evidence for direct uptake of fog condensation on leaves by plants. Any leaf structure
capable of absorbing water on the leaves is also a potential route via which water can
evaporate. Thus, there would be little benefit for plants in a hyper-arid environment to absorb
fog moisture directly from the leaves. A more likely route whereby plants could benefit from
fog moisture would be by absorbing condensation on the sand surface and as a result of stem
flow. This route would facilitate the uptake of both fog and dew condensation on the soil
surface. Certain plants growing in the fog zone of the Namib have well-developed superficial
root networks or efficient mycorrhizal relationships to be able to benefit from alternative
moisture sources such as fog and dew. In the Namib Desert lichens grow in great diversity on
west facing slopes and surfaces where they are able to draw moisture from the sea fogs. If it
were not for the fog the plants would have no source of water (Lange 2003). Dawson (1998)
has clearly documented the importance of fog drip for soil moisture and root uptake. In the
Arctic, foggy weather cut down the photosynthetic rates, but the net amount of carbon fixed
by the end of growing season is high at about 174 g C per m2 per season (Billings and
Peterson 1992). Gas exchange is completed by small pores in the leaf surface called stomata.
Most desert plants have few stomata on the bottom side of the leaf, protected by deep
depressions. The relationship between fog and lichen flora in the Antarctica is one of the good
examples.
Fog is important as a source of water for organisms, which cannot take up water from the
substrate, such as lichens on rocks. It is also important for lichens to be able to
photosynthesize continuously in the daytime, which requires sufficient light and moist (but
not wet) conditions. Fog here is more favourable to lichens than rain, i.e. especially fog in the
daytime. In climates with mainly night fog and drought during the day lichens do not thrive,
because respiration at night continues and is not compensated for sufficiently by
photosynthesis (a so-called negative photosynthetic balance). In the SO, for example, the
number of days with fog is generally low throughout the years. In the same direction the
epiphytic lichen flora is considerably impoverished. Thus, maps of fog frequency are
important if one wants to know whether the fog habitually occurs at night or in the daytime
and affects lichen flora (Gajananda et al., 2007). In SO the solar heating of the ground and
formation of thermal plumes had less effect on the formation and dissipation of the fog
episode. The dissipation of fog was only after the katabatic wind started flowing from the
interior of the continent. Therefore, this type of fog can be regarded as advection fog (figure
10). It is also likely that due to changing climate over Antarctica, frequent fog may occur in
the near future. The monostatic acoustic sounding data in association with the study of
microbiotic crusts retention of atmospheric moisture especially during foggy period will help
in understanding the complex physiological and metabolic characters of the various
microbiota of Antarctica. The cytological mechanisms of the microbiotic communities of SO,
which used fog moisture are unclear. Therefore, research on this aspect will be fruitful.
78
Figure 11. Facsimile chart showing thermal plumes on January 10, 1996 at Maitri. It shows plumes
around 0945 to 2200 hrs.
79
Short spells of winter warming enhancing surface air temperature by even 15 0C have
been measured. Such warming events may play an important role in the sustenance of
microorganisms from totally desiccating their body fluids (Gajananda 2004b). These
convective or unstable atmospheric conditions in the lower atmospheric boundary layer may
help carry upward the spores/pollens of the micro-flora in the SO and facilitate their
dispersive capacity to a larger distance (Gajananda et al., 2004).
4. CONCLUSION
From the above studied, it is clear that Antarctica provides a unique platform for
understanding the climatic processes in a complex environment. Study of land-ice-air-ocean
interaction over the SO reveals that a unique ecosystem prevails over this oasis, where all the
physical and biological components interact in a complex pattern.
The conclusions drawn from the present research are summarized as:
1. The climate at Maitri is dominated by the extreme contrasts between the seasonal
inputs of solar radiation. It experiences sub-zero mean temperature throughout the
year except in the peak summer months (December and January). The surface wind
regime (annual mean 10.2 m/s) is dominated by katabatic wind component from the
southeast direction (110 to 1800), which is the direction of maximum slope around
the Maitri site. Pressure (annual average 986.5 mb) forms half yearly cycle and
influenced for the formation of cyclones. The humidity and precipitation are although
low but have significant relationship for the growth of microorganisms. Surface
albedo also has great effect on the warming and cooling of the surface of SO and in
formation of atmospheric convection.
2. The monthly average surface air temperature observed at Novolazarevskaya station
shows that the temperature has significantly increased during the last four decades by
1.11 0C (P<0.01), which is at the rate of about 0.27 0C per decade. The summer
(December-February) of this period also shows a significant raise in temperature by
about 0.99 0C (P=0.05), which is at the rate of about 0.24 per decade increase. Hence,
in the SO the surface air temperature enhancement will certainly affect the function
of ecosystem dynamics.
3. The ecosystem of SO must have grown with some control of high UV-B doses. The
threshold limit of UV-B (1 MED/hr) was crossed during both spring and summer for
the study period. Thus, a profound effect on the flora and fauna is likely. A
comprehensive long term program involving the study of ecosystem under both
prolonged low UV-B conditions and high UV-B conditions should be drawn, so as to
understand the relationship and scientific mechanisms through which the ecosystem
of SO has survived.
4. Thick snow cover forms an important part of the climate system and affects the
ecology of the region. The snow acts as an energy bank by storing latent heat of
fusion and sublimation and crystal bonding forces. The intake and release of energy
throughout the year seems to be responsible for the variability in habitat conditions
leading to heterogeneity of the ecosystem composition at the SO. The snow has a
80
It is clear from the above points that all the climatic parameters are limiting the survival
of living organisms in SO. The floral diversity in SO is poor and dominated by primitive
poikilohydric groups of producers (algae, mosses and lichens) forming thin crusts on the soils
surface. It also exhibits a very narrow spectrum of micro-floras in two distinct ecosystems i.e.
terrestrial and aquatic. With the abiotic factors limiting life forms, the following conclusions
are made:
1. The energy cycle and food chain are poor and short in SO due to less sunlight during
austral winter and lack of higher organisms.
2. About 34 species of primitive flora were observed and only 6 primitive invertebrate
fauna were recorded. Only four species of birds were observed in SO. The algal
species are dominated by blue green algae (cynobacteria) and green algae are not
frequent. Lichens are the most wide spread species in this oasis. Mosses have less
species diversity but their frequency over the oasis is large.
81
ACKNOWLEDGMENTS
We are grateful to the Director, National Physical Laboratory (NPL), New Delhi and
to the Secretary, Ministry of Earth Sciences, Government of India for providing us an
opportunity to work at the Indian Antarctic station, Maitri. We also thank the Director (Dr.
Rasik Ravindra), National Centre for Antarctic and Ocean Research (NCAOR), Goa for the
supports and logistics helps. The constructive criticisms rendered by Professor Anubha
Kaushik, Guru Jambheshwar University, Hisar is thankfully acknowledged. The Leader (Dr.
Ajay Dhar) and Members of the 18th and 19th Indian Scientific Expedition to Antarctica
(InSEA) who helped in surveying the SO is thankfully acknowledged. Thanks are also due to
IMD at Antarctica and to Dr. Risal Singh and Mr. Shambhu Nath of NPL, New Delhi who
provided data of the harsh polar environment. The financial assistance to one of the author
(KG) under CSIR-SRF, DOD-SRF (DOD/12-MMDP/897-MRC CELL) and deputation as a
Wintering Member of the 18th InSEA (19982000) by NPL, New Delhi, is acknowledged
with thanks.
82
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Chapter 4
90
INTRODUCTION
Till 1830, Antarctica remains as an unsolved puzzle for researchers. Later on number of
expeditions to this continent were carried out by the explorers and scientists, from different
countries basically for collecting flora and fauna from the different parts of the icy continent.
In the year, 1948 the British, Australian, New Zealand, conducted Antarctic research
expeditions for exploring the biodiversity of flora along with meteorological analysis at
Antarctica. Antarctica is almost entirely (99.68 %) covered by permanent ice and has a
surface area of 13,830,000 sq. km., which include several massive floating ice-shelves, which
is some what similar to the area of Arctic Ocean (British Antarctic Survey 2004). Three major
national voyages were carried out the Antarctica between 1837 and 1843; an important object
for each was the determination of the position of the South Magnetic pole. The French naval
expedition (1837-1840) visited the South Orkney Islands, South Shetlands, Trinity peninsula,
Orleans Strait and Joinville Island. The International Polar Commission was formed at a
conference in Hamberg, 1889 and a second conference was held in Berne, 1880 and a third in
Sankt Peterburg, 1881, (Headland 2005).
The international polar year conducted from 1st August 1882 to 31st August 1883, and 12
countries established 14 scientific stations in polar regions for co-coordinated Scientific
observations. Three national expeditions, from Britain, Germany and Sweden set out in 1901
after much careful planning, the intended to explore different part of Antarctica and coordinate their geophysical and meteorological observations (Headland 2005).
Indias interest in Antarctica arises mainly from its Gondwana land connection, the
lack of any land barrier between the Indian and Antarctic Ocean is also of great significance.
Indias first research station Dakshin Gangotri commissioned in the year 1984 and second
research station Maitri commissioned in the year 1988-89 over the exposed rocks of
Schirmacher Oasis of Queen Maud Land area in east Antarctica (Figure 1). The Schirmacher
Oasis (700 4558S) is one of the small, snow / ice covered during the winter (June-August)
and spring (September-November) seasons, typical desert near the shore of East Antarctica,
with a width of 1.6 Km and length of about 20 Km with east-west orientation.
91
Its composed of with mainly high grade of metamorphic rocks, the fresh water lakes,
ponds, pools cover a total area of 35 square km, and the altitude varies from 0 to 228m with
an undulated surface.
Climatic conditions are similar to other East Antarctica ice-free areas with annual
temperature of -100C, January is the warmest month (-0.60C) and August is the coldest (10.10C), (Francisco et al. 2004). Recently India started to build its third station at Larsemann
Hills, (69o20S to 69o30S Lat.; 75o55E to76o30E. long) in Prydz Bay, is an ice-free oasis on
the Ingrid Christensen coast, of East Antarctica.
92
27
Lichens
Umbilicaria, decussata
Acarospora gwynnii
Buellia pallida
Lecidea cancriformis
Lecanora fuscobrunnea
Physcia caesia
Rhizocarpon flavum
Rinodina olivaceobrumea
Umbilicaria antarctica
Xanthoria elegans
Usnea antarctica
Sphaerophorus globosus
Parmelia saxatilis
Turgidosum complicatulum
Pseudephebe minisciila
Rizoplaca melanopthalma
Carhonea capsulata
Pseudephebe miniscula
Bryophytes
Bryum argenteum
Bryum pseudotriquetrum
Ceratodon perpureus
Grimmia antarctici
Sanionia uncinata
Lichens
Andrea regularis
Cephaloziella varians
Schistidium antarctici
Algae
Entomeneis kjellmannii
28
Berkeleya adeliense
29
Nitzschia stellata
30
31
Prasiola crispa
Choricystis minor
32
33
34
35
36
Phormidium sp
Lyngbyasp sp
Oscillatoria sp
Nodularia sp
Anabaena sp
19
20
21
22
23
SN
24
25
26
Location
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Shirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Antarctic Peninsula
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
References
Czeczuga 1996
Dodge 1973
Dodge 1973
Dodge 1973
Dodge 1973
Czeczuga 1996
Dodge 1973
Dodge 1973
Lamb 1964
Czeczuga et al. 1986b
Lamb 1964
Czeczuga et al. 1986b
Czeczuga 1996
Lud et al. 2001
Upreti et al. 2000
Upreti et al. 2000
Upreti et al. 2000
Upreti et al. 2000
Antarctic Peninsula
Windmill Islands
Windmill Islands
Windmill Islands
Antarctic Peninsula
Location
Antarctic Peninsula
Antarctic Peninsula
Windmill Islands
Lichens
Chlorell vulgaris
Chlorococcum sp
Phormidium uncinatum
Oscillatoria sp
Aphanothece sp
Urospora sp
Cylindrocapsa geminella
Calothrix brevissuna
Chrococcus macrococcus
Gloeocapsa sp
Lyngbya martensiana
Nostoc commune
Stigonema sansabaricus
Higher Plants
Deschampsia antarctica
Colobanthus quitensis
Location
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
Schirmacher Oasis
References
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Shukla et al. 1999
Antarctic Peninsula
Antarctic Peninsula
Lichens
Umbilicaria,
decussata
Acarospora
gwynnii
Buellia pallida
2
3
4
5
6
7
8
9
10
11
Lecidea
cacriformis
Lecanora
fuscobrunnea
Physcia caesia
Rhizocarpon
flavum
Rinodina
olivaceobrumea
Umbilicaria
antarctica
Xanthoria
elegans
Usnea antarctica
Photosynthetic
Pigments
Other pigments
References
-Carotene, Lutein,
Cathaxanthin, Astaxanthin
-Carotene, Lutein,
Hydroxyechinenone
-Carotene, Lutein,
Zeaxanthin
-Carotene, Lutein,
Czeczuga 1996
-Carotene, -Carotene,
Lutein
-Carotene, Cryptoxanthin, Lutein,
Astaxanyhin
-Cryptoxanthin,
Canthaxanthin, Lutein,
-Carotene, Lutein,
Zeaxanthin, Canthaxanthin
-Carotene, Cryptoxanthin, Lutein,
-Carotene, Cryptoxanthin, Lutein,
Asthaxanthin
-Carotene, Cryptoxanthin, Lutein,
Hydroxyechinenone
Czeczuga, 1996
Czeczuga 1996
Czeczuga 1996
Czeczuga 1996
Czeczuga 1996
Czeczuga 1996
Czeczuga 1996
Czeczuga 1996
Czeczuga 1996
Czeczuga 1996
93
94
Lichens
12
Sphaerophorus
globosus
Parmelia
saxatilis
Turgidosum
coplicatulum
Bryophytes
Bryum argenteum
Other pigments
References
Astaxanthin, Adonixanthin
Czeczuga 1996
Czeczuga 1996
Violaxanthin, - Carotein
Bryum
pseudotriquetrum
Ceratodon
perpureus
Grimmia
antarctici
Violaxanthin, - Carotein
Violaxanthin, - Carotein
Violaxanthin, - Carotein
Sanionia
uncinata
Andrea regularis
25
Cephaloziella
varians
Schistidium
antarctici
Algae
Entomeneis
kjellmannii
Berkeleya
adeliense
Nitzschia stellata
Neoxanthin, Lutein,
Zeaxanthin, ,- Carotene
Neoxanthin, Violaxanthin,
Lutein, Zeaxanthin
Neoxanthin, Lutein,
Zeaxanthin, ,- Carotene
Anthocyanin
Markham et al.
1990
Robinson et al.
2002; 2006
Dunn 2000
Robinson 2002
Dunn 2000
Robinson 2002;
2005
Newsham et al.
2002
Newsham 2003
26
Prasiola crispa
27
28
Choricystis minor
Phormidium sp
29
Lyngbya sp
Chl a.,
30
Oscillatoria sp
Chl a.,
31
Nodularia sp
Chl a.,
13
14
15
16
17
18
19
20
21
22
23
24
Photosynthetic
Pigments
-carotene
MAAs (porphyra-334) Carotene
MAAs (porphyra-334) Carotene
MAAs (porphyra-334) Carotene
MAAs (porphyra-334) Carotene
Newsham et al.
2002; 2005
Robinson et al.
2006
Ryan et al.
2002
Ryan et al.
2002
Ryan et al.
2002
Post and
Larkum 1993
Zidarova 2006
Quesada et al.
1998
Quesada et al.
1998
Quesada et al.
1998
Quesada et al.
1998
Lichens
32
Anabaena sp
33
Palmaria
decipiens
Phyllophora
antarctica
Porphyra
endiviifolium
Iridaea cordata
34
35
36
37
38
Higher Plants
Deschampsia
antarctica
Colobanthus
quitensis
Photosynthetic
Pigments
Chl a.,
Other pigments
References
Quesada et al.
1998
Dhargalkar
2004
Dhargalkar
2004
Dhargalkar
2004
Dhargalkar
2004
Orientin, Luteolin
Orientin, Luteolin
Ruhland et al.
2001
Ruhland et al.
2001; Lud et al.
2001
Chl a.,
Chl a.,
Chl a.,
Chl a.,
95
96
may be helpful to accumulate high photo energy from sun and retention in lower amount,
these melanin help to protect lichen thallus against UV-radiation. Gauslaa and Ustvedt,
(2003) explored the brightly orange color anthroquinone i. e. parietin found in lichens and
discussed their role as UV screening compound and concluded that parietin may reduce the
effect of UV radiations. UV-B absorbing pigments such as flavonoids are wavelength
selective UV-B screens, which can accumulate rapidly in response to high UV-B radiation
levels, Caldwell et al. (1983). Morphological changes includes reduced growth, where as
reduced photosynthesis is also important parameter for measurement (Prasad et al. 2005).
Several pigments like chlorophyll, carotenoid in lichens, algae, bryophytes, and green alga are
the advance parameter rather than morphology and anatomy, to detect the effect of UV
radiation on Antarctic flora. The pigment found, in photoautotrophic organisms have the
interest to ecologist and biologist for a number of reasons, e. g. chlorophyll and their
derivatives has been much used a parameter for the measurement of productivity. Their
qualitative as well as quantitative measurements may be indicative of the climatic changes.
LICHENS
The peculiarity of lichens is that they are not one homogenous organism but a symbiosis
of two different partners. The fungal partner supplies the plant, water and nutritious salt,
meanwhile the algal partner supplies the organic substances, like carbohydrate produce. With
this ideal job-sharing lichens can withstand hardest environmental conditions. Of all the
plants, lichens are best adapted to survive in the harsh polar climatic conditions (Table 1).
Some lichens have even been found only about 400 km from the South Pole. Lichens are
colorful organisms owing to numerous combinations of algal and fungal pigments; color is
often used in taxonomic studies to aid species identification. Some times the color of the
thallus is used as a characteristic to discriminate related genera; however, color variation
within species can be significant because of spatial and temporal variations through
environmental factors (Gauslaa and Mc Evoy 2005). Among Antarctic flora, the lichen shows
extensive diversity, thus it has taxonomical, ecological, environmental, interest of many
lichenologist from various countries. According to Dodge (1973) there are 429 species of
lichens occurring in Antarctica, while Hertel (1988) estimated only 160 crustose lichens in
that area. Comparing the investigation to Indian context a single lichen genus Acarospora was
recorded by Wafar and Untawale (1983), from Maitri area. Upreti (1999) collected lichens
from Maitri region of Schirmacher Oasis, and enumerated about 23 species of lichens
belonging to 16 genera. According to Friedmann (1982), in Antarctica, lichens are the major
floral elements of terrestrial vegetation of the harsh climate, after the disappearance of macrolichens the rock inhabiting (saxicolous) crustose species forms the main component of the
lichen vegetation. The crustose lichen develops a thin primitively organized thallus, which is
completely affixed to the substratum over the whole lower surface or hidden in the uppermost
layer of the rock. According to Hertel (1988), the crustose lichens can colonize, in extreme
polar habitat better than the other macroscopically visible plants. The common genera of
lichens, are reported in Table 1 are Acarospora, Buellia, Candellariella, Lecidea, Lecanora,
Physcia, Rhizocarpon, Rinodina, Umbilicaria, Xanthoria, Turgidosculum, Usnia, etc are
luxuriant growing in Antarctica (Czeczuga 1996; Lud et al. 2001; Lamb 1964; Dodge 1973).
97
The poikilohydric nature of lichens and several secondary chemicals like atronorin, usnic
acid, zeorin, triterpins are widely found, and help to protect against stress; thallus color of
lichens i. e. orange, red, yellow, grey are also helpful during harsh environmental conditions.
During stress conditions these secondary chemicals reach at upper cortex of lichens thallus to
protect them (Elix, 1996). Leading to these types of studies Huneck (1999), justified that
these secondary chemicals some time react with rock substratum and secrete powdery
substance on upper surface called Pruna these pruna shade the thallus to protect the thallus
against temperature as well UV radiations. The darkness of melanin pigments are observed
with the naked eye in intact Lobaria pulmonaria thallus again a natural light gradient reflects
the level of solar exposure. The brownest thalli are observed in the most sun-exposed position
and increasing pigmentation causes a reduced cortical transmittance, especially of UV and
short wave visible radiation. A photo protective role for melanin compound should imply a
higher resistance against excessive light in thalli where these pigments are abundant (Riley
1997). The melanic green algal lichens like Parmelia and Cetraria display predominantly
Arctic / Alpine regions; this melanin shows the strongest protection at energy-rich
wavelength, the percentage reduction in transmittance caused by melanin appears to be large
in the UV range. The secondary medullary compounds, stictic and norstictic acid deposited in
largest quantities in the photobiont layer may assist in UV protection (Bachereau and Asta
1997).
BRYOPHYTES
Like the lichens, the bryophytes can be found in all areas capable of supporting plant life
in the Antarctic, though they are not very widespread in distribution. Bryophytes, even those
of open, exposed habitat have been seen as showing features that would generally be regarded
as characteristic of shade plants. With lichens, bryophytes share the distinction of being the
largest group of plants that occur in Antarctic region. In spite of general assumption that
Joseph D. Hooker made the first collections of mosses in Antarctica during 1829-30. The
bryophytes are also poikilohydric, depending on the presence of free water during the summer
month for photosynthetic carbon gain and growth. Antarctic bryophyte communities are
largely confined to the margins of melt lake at Casey (the Australian base in the Windmill
Island region, 660 17S, 1100 32E) the three dominant moss species Ceratodon purpureus,
Grimmia antarctici and Bryum pseudotriquetrum are found in the both pure and mixed
communities (Selkirk and Seppelt 1987) and stream areas subject to snow accumulation. In
addition to their role in energy storage compounds, simple sugars and polyols performs
multiple roles in plant. They are thought to act as cryoprotctants, as osmotic regulator in the
drought and salt stress plants, and as antioxidant (Popp and Smirnoff 1995). Various soluble
carbohydrate are able to interact with the polar head groups of phospholipids, taking the place
of water molecules and maintaining membrane integrity during desiccation (Crowe and
Crowe 1986), polyols accumulate in higher plants in response to water stress. These
molecules probably have multiple roles, as compatible solutes, scavengers of active oxygen
species and as stabilizers of macromolecules (Loescher 1987; Smirnoff and Cumbes 1989;
Popp and Smirnoff 1995). Differences in the response of the three moss species to
desiccation, and the phenotypic plasticity in their response, was assessed by testing for
98
differences in three parameters over two sites that differed in water availability; (1) the WC
(water content) at full hydration, (2) the rates of drying over time, and (3) the relationship
between the decline in chlorophyll fluorescence and relative WC. Initial rates of water loss
were faster in moss obtained from the wet sites compared to that from the dry sites.
Few mosses have been studied, to know the negative effects of UV radiation on growth
and morphology in some high latitude species (Sonesson et al. 1996; Searles et al. 1999;
2002, Robson et al. 2003, Robinson et al. 2005). Short moss turf and cushion moss is found
most frequently in sandy and gravelly soil. The extent of these moss communities depends on
the availability of summer melt water, and ranges from the extensive moss which occurs
around, melt lakes and along melt streams (Selkirk and Seppelt 1987). Mosses, like lichens,
gather in colonies, which make them possible to collect and retain more water. They also lose
less by evaporation and show a marked ability to use water rapidly whenever it becomes
available. Mosses have also become well adapted to the almost continuous light during long
day of a polar summer. The impact of ambient ultraviolet (UV-B) radiation on the endemic
bryophyte, Grimmia antarctici, was studied over 14 months in East Antarctica. Over recent
decades, Antarctic plants have been exposed to the largest relative increase in UV-B exposure
as a result of ozone depletion. Although photosynthetic rates were not affected, there was
evidence of UV effects on morphology Antarctic bryophytes possess UV-B absorbing
pigments which should offer better protection under ambient UV-B radiation, these findings
suggested that G. antarctici may be disadvantaged in some settings under a climate with
continuing high levels of springtime UV-B radiation (Robinson et al. 2005).
Recently, high concentrations of UV-B absorbing pigments have also been reported in
two Antarctic mosses, Sanionia uncinata and Andreaea regularis, and one liverwort,
Cephaloziella varians, with positive correlations between pigment accumulation and flux of
natural solar UV-B radiation (Newsham et al. 2002, 2005, Newsham 2003), and investigated
the effect of reduced UV and visible radiation on the pigment concentrations, surface
reflectance and physiological and morphological parameters of the moss. Chlorophyll content
was significantly lower in plants grown under near-ambient UV, while the relative
proportions of photoprotective carotenoids, especially -carotene and zeaxanthin, increased
(Table 2). However, no evidence for the accumulation of UV-B absorbing pigments in
response to UV radiation was observed. According to Paul (2001), the carotenoid
concentration plays an important role in prolonged period of freezing. The Bryum argenteum
produce more energy by photosynthesis in low light at 50C than it does at 150C or higher.
Photosynthesis can start within few hours of thawing after a prolonged period of freezing, and
almost immediately following short period, due to this reasons it may be survive there.
ALGAE
Snow algae, which grows in semi permanent to permanent snow or ice in the alpine or
polar region of the world. Their optimum growth temperature is generally below 100C.
According to Green et al. (1999) more than 300 species of non-marine alga have been found
in Antarctica, these algae successfully adapted to their harsh environment though the
development of a number of adaptive features which includes pigments polyols (sugar
99
alcohols, e. g. glycerin), sugar and lipid (oils), mucilage sheaths, motile stages and spore
formation.
Cyanobacteria (blue green algae) have a cosmopolitan distribution ranging from hot
springs to Arctic and Antarctic regions characterized by high variability and the organisms
need to adapt to various environmental factors (Rozema et al. 2002). Blue green and other
algae are found growing in damp sand and gravel around lakes and pools along melt water
streams or in low-lying areas.
Other than blue and green algae may form extensive and spectacular red, yellow or green
patches in area of permanent snow. These pigments protect the cells from the high light and
ultraviolet radiation damage during the summer months. The pigments may take the form of
iron tannin compounds, as M. berggrenii, or orange to red-pigmented lipids as in the majority
of the snow algae. The cells of some species also secrete copious amount of mucilage, which
unable them to adhere to one another and to snow crystal and to prevent the cell from being
wash away by melt water. The mucilage also forms a protective cover and delays desiccation.
It may have additional function as an ultraviolet shield. According to Sinha et al. (2005),
Ultraviolet-B radiation reaching the earth surface due to depletion of stratospheric ozone layer
have been shown to cause destruction of phycobiliproteins like phycoerythrin, phycocyanin,
allophycocyanin, especially in blue green algae. In general, cyanobacteria are protected by
mycosporine-like amino acids (MAAs) and scytonemins while terrestrial plants contain
flavonoids (Table 2).
FUNGI
In Antarctic conditions, fungi have been studied in little. Several mushrooms have been
found on the west coast of the Antarctic peninsula, and on the South Shetlands. A few of the
fungi in Antarctica are unique to the continent and responsible to deteriorate the quality of
material but there is little study regarding pigments (Green et al. 1999).
HIGHER PLANTS
Only two native vascular plants, the Antarctic hair grass Deschampsia antarctica and a
cushion forming pearlwort, Colobanthus quitensis, survive south of 560 S. They occur in
small clumps near the shore of the west coast of Antarctic Peninsula. Both plants can tolerate
very harsh environmental conditions. The accumulation of UV-B absorbing pigments would
be particularly useful in Antarctic plants because such passive screens could protect them
from UV-B damage when physiological inactivity, due to desiccation or freezing, renders
active repair mechanisms unavailable (Lovelock et al. 1995, Cockell and Knowland 1999).
They continue to maintain their physiology at freezing point (Caldwell et al. 1998).
100
induce lipid and protein peroxidation and thus, these active oxygen species are suggested to
affect the structural integrity and permeability of cellular mechanisms due to ultraviolet
radiations. Almong et al. (1991) suggested that enhanced UV-B radiation causes a multitude
of differing effects such as change in electron transport capability, chloroplast ultrastructure,
level of photosynthetic and protective pigments and disturb the mechanism of photosynthesis
in terrestrial plants. Among two photosystem, photosystem II appeared to be more sensitive to
ultraviolet radiation than photosystem I. Significant reduction in chlorophyll content was
noticed with increasing dose of UV-B and the suppression was more prominent on
chlorophyll-a than chlorophyll-b in other words it shows decreasing trend in Chl a / Chl b
ratio. This reduction could be due to chlorophyll destruction as reported in most of the UV-B
exposed photosynthetic organisms in glass house study (Teramura and Sullivan 1994). In
contrast to chlorophyll contents, carotenoid contents showed an enhancement with increasing
dose of UV-B radiation. In order to avoid AOS induced damage during oxidative stress,
plants evolve a complex antioxidant defense system (Kondo and Kawashima 2000). The
antioxidative enzymes: superoxide dismutase, catalase, peroxidase, etc. and non-enzymatic
antioxidants: ascorbate, carotenoids, flavonoids and a variety of phenolics are synthesized in
stressed plants (Rao et al. 1996). UV-B may cause damage to nucleic acids, proteins, and
lipids, and may affect photosynthesis, growth and development of plants (Jansen et al. 1998).
According to Tevini et al. (1981), carotenoid acts as scavenger of singlet oxygen which is
formed during intense light, protect chlorophyll from photo-oxidative damage therefore,
increased carotenoid content following UV-B exposure could have a positive effect on
protecting chlorophyll pigments. Interestingly, concentrations of UV-B absorbing compounds
were higher and chlorophyll-b was lower under ambient UV-B radiation in two Antarctic
vascular species, D. antarctica and C. quitensis, (Ruhland and Day 2000). There is also
evidence to suggest that reductions in leaf area may be related to increases in insoluble levels
of UV-B absorbing phenylpropanoids that contain epidermal cell elongation and subsequent
leaf expansion. Studies have found that the ambient levels of UV-B can stimulate production
of UV-B absorbing compounds (Hunt and Mc Neil 1999).
It is important to study the effect of particular stress on the biological organisms, and to
explore the pathway for signaling system i. e. defense mechanism at targeted organism and
also explore the mechanism between photosynthetic pigments and defense. Among these it is
also necessary that what type of defense be adopted by the organisms during adverse
conditions. In Antarctica, there are intense ultraviolet radiations therefore, it is important to
study the effect of ultraviolet radiation on flora and fauna and type of defense adopted for
surviving organisms. Leading to these types of studies in Robson et al. (2003) using polyester
film for the attenuation of ultraviolet radiation, which transmit a large percentage of the UVA radiation and found that photosynthetic pigment altered with UV-B radiations. Phoenix et
al. (2002), have used polycarbonate filter that absorb both UV-A and UV-B radiation, in this
experimental model the effect of ultraviolet radiation, in subarctic shrubs, reductions in UV-B
radiation resulted in increased levels of UV-B absorbing compounds which was attributed to
either increased secondary metabolism in absence of UV-B radiation or UV photo-oxidation
of these pigments. Krizek and Mirceki (2004) reported that cellulose acetate film as sometime
used for near-ambient control in UV-B exclusion or UV-B attenuation experiments can have
toxic effects on plants and affect growth in dependently of their effect on UV radiation. The
experiment of Robson et al. (2003) is not affected by this problem as Aclar film was used
instead of cellulose acetate for in-vitro as well field study. Result of these UV-B experiments,
101
REFERENCES
Almong O, Lalan O, Shoham G, and Nechushtai R, (1991) The composition and organization
of photosynthesis I. J. Basic Clin. Physiol. Pharmacol. 2: 123-140.
Barsig M, Malz R, (2000) Fine structure, carbohydrates and photosynthetic pigments of suger
maize levels under UV-B radiation. Environ. Exper. Bot. 43: 121-130.
Bachereau F, Asta J, (1997) Effects of solar UVR at high altitude on the physiology and the
biochemistry of terrestrial lichen Cetraria islandica (L.) Ach. Symbiosis 23: 197-217.
Blumthaler M, and Ambach W, (1990) Indication of increasing solar UV-B radiation flux in
alpine regions. Science 248: 206-208.
Bothwell M L, Sherbot D M T, and Pollock C M, (1994) Ecosystem response to solar
ultravioletB radiation: Influence of trophic-level interactions. Science 265: 97-100.
British Antarctic survey, (2004) Antarctica (map at 1:10,000,000 scale). Miscellaneous
publication 11: Cambridge, British Antarctic Survey.
Caldwell M M, Flint S D, Searles P S, (1994) Spectral balance and UV-B sensitivity of Soya
bean: a field experiment. Plant Cell Environment 17: 267-276.
Caldwell M M, Bjorn L O, Bornman J F, Flint S D, Kulandaivelu G, Teramura A H, and
Tevini M, (1998) Effect of increased solar UVR on terrestrial ecosystems. J. Photochem.
Photobiol. B. Biol. 46: 40-52.
Caldwell M, Robberecht R, Flint S, (1983) Internal filters: prospects for UV-acclimation in
higher plants. Physiologia Plantarum 58: 445-450.
Cockell C S, Knowland J, (1999) Ultraviolet radiation screening compounds. Biological
Review 74: 311-345.
Crittenden P D, (1998) Nutrient exchange in an Antarctic macro-lichens during summer
snowfall-snowmelt events. New Phytologist 139: 697-707.
Crowe J H, Crowe L M, (1986) Stabilization of membranes in anhydrobiotic organisms. In
Membranes, metabolism and dry organisms Cornell University Press: Ithaca, USA (Ed.
C Leopold) pp 188230.
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103
104
105
106
Chapter 5
1. INTRODUCTION
Antarctica is a place like no other; as an intriguing habitat, it is a scientist's dream. It is a
land where water is scarce, despite having more than two-thirds of the world's freshwater
supply trapped in ice. Though it borders the world's major oceans, the Southern Ocean system
is unique; it is a sea where average temperatures do not reach 2C in summer, where even the
water is so unusual that it can be identified thousands of kilometers away in currents that
*
Email: jaswant1983@yahoo.co.in
Department of Environmental Sciences, Dr. R.M.L. Avadh University,
Faizabad-224001, U.P., India
108
originated here. As the Earth, tilt on its rotational axis, makes its elliptical journey around the
Sun each year, the Sun "sets" in April, not to be seen again until September. And the icean
unimaginable, incomparable vastness of iceappears in a dozen different varieties, at times
and in places several thousand meters thick. There are two major ice sheets that change all the
time. Antarctica is a remote and inhospitable continent. The climate is the coldest and driest
known on Earth; nevertheless it is not uniform across the continent, and different climatic
regions can be distinguished (Holdgate 1977; Onofri 1999; vstedal).
The prevailing Antarctic conditions of low temperature, low water availability, frequent
freezethaw cycles, low annual precipitation, strong winds, high sublimation and evaporation,
high incidence of solar and especially ultraviolet radiation together constitute significant
limiting factors for plant and animal life. As a consequence of these severe conditions, the
Antarctic flora is almost entirely cryptogamic, only two vascular species occur, both of which
are restricted to the relatively mild Antarctic Peninsula. Therefore, the biology of Antarctica,
more than other continents, is dominated by microorganisms, with a high level of adaptation
and able to withstand extreme conditions. Abundance and diversity of organisms decrease,
along broad latitudinal gradients, from the maritime to the continental Antarctic zone and
within the latter with increasing altitude and latitude from the coast to the Ice Slope Region
(Pickard and Seppelt 1984; Kappen 1993).
109
ranges from the extensive moss turves which occur around melt lakes and along melt streams
to small isolated moss buttons which are found in moisture pockets on rocky outcrops. Of the
three species, G. antarctici is common within melt lakes and streams, C. purpureus is often
associated with drier sites, and B. pseudotriquetrum co-occurs with both G. antarctici and C.
purpureus (Selkirk and Seppelt 1987) Despite limitations to growth , mosses have been
reported from as far south as 84.1 S and although bryophyte fruiting events are rare in the
continental Antarctic zone (Wise and Gressitt, 1965; Filson and Willis, 1975). In addition to
the relatively conspicuous mosses and lichens, the continental Antarctic terrestrial vegetation
includes groups that are often overlooked, including the chasmoendolithic algae, which occur
only within rock fissures. These organisms are widespread in coastal regions of Antarctica
and are believed to underlie up to 20% of the rock surface in some locations (Longton, 1985;
Hansom and Gordon, 1998). Total plant group contributing to the biodiversity of the
Antarctic landmass is provided in the table 1.
Table. 1. Estimated contribution of plant groups to the terrestrial plant
biodiversity of Antarctica
Phytogeographic
zone
Continental
Maritime
Total
Angiosperms
Mosses
2
2
30
75
85
Liverwort
s
1
25
26
Lichens
125
150
200+
Macrofung
i
2
22+
28
References
Lewis Smith(1984)
Lewis Smith(1984)
Longton(1985)
Figure 1. Map of Antarctica. The Maritime Antarctic is the area to the left of the dashed line. The
Periantarctic islands are found north of the limit of sea ice and bounded by the polar front. Locations
where much of the research described in this review was conducted include Signy Island, Maritime
Antarctic, Windmill Islands, Wilkes Land, Ross Island and Victoria Land ( Adapted from Robinson
2003).
110
Figure 2. Ozone hole area w.r.t to 220DU in Southern hemisphere from year 1999 to 2008 (NASA
2008).
111
Figure 3. Effects of UV-B radiation on plant cells, showing screening sites of damage.
112
The annual occurrence of the ozone hole coincides with time of emergence from winter
dormancy beneath the protective snow cover (Karentz, 1991; Adamson and Adamson, 1992;
Wynn-Williams, 1994), exposing plants to sudden elevations of UV-B radiation in
combination with increased PAR and greater temperature fluctuations. Bryophytes may be
particularly susceptible to UV-B damage because of their simple structure, with most having
leaves that are only one cell thick and lacking protective cuticles or epidermal layers
(Richardson, 1981; Gehrke, 1998; Gwynn-Jones et al., 1999). Combined with the
physiologically stressful effects of repeated freeze/thaw cycles, an intermittent water supply
and limiting nutrients, polar bryophytes are likely to be sensitive to the additional stress
imposed by elevated UV-B radiation (Robinson et al., 2003;Wasley et al., 2006 a, b). The
survival of Antarctic plants under ozone depletion depends on their ability to acclimate, by
employing photoprotective mechanisms to avoid and repair UV-B damage.
Many of the effects induced by UV-B radiation allow for at least a duality of
interpretations. The difference between damage, repair and acclimation can be subtle and it is
not always possible to identify one particular mechanism as the explanation underlying a
given phenomenon. For example, the UV-B induced degradation of the D1 protein of PSII
can be seen either as damage or as a part of a repair mechanism leading to the substitution of
the damaged components of PSII. A number of studies have found that photosynthetic
activity (estimated by measuring gas exchange or chlorophyll fluorescence) does not appear
to be strongly affected by either reduced or elevated UV-B treatments in the two Antarctic
vascular species. However, growth was affected in a number of ways by exposure to UV-B.
The major impact was a reduction in cell length, leading to shorter leaves. Less branching and
fewer leaves per shoot led to reduced plant size and biomass, with effects more pronounced in
C. quitensis than D. antarctica. Leaves were also thicker in plants exposed to UV-B. Perhaps
the cost of producing and maintaining thicker leaves explains the reductions in growth that
occur in the absence of effects on photosynthesis (Xiong and Day, 2001; Xiong et al., 2002).
Long-term field studies showed similar but less pronounced impacts of UV-B radiation
compared with similar length pot studies, but the latter showed high inter-annual variation
and provided evidence of cumulative UV-B effects (Day et al., 2001). Exposure to UV-B
accelerated plant development and led to greater numbers of reproductive structures in both
species, although the weight of C. quitensis seed capsules declined with higher UV-B
exposure. However, since these structures produced fewer spikelets and seeds, the overall
reproductive effort was unchanged. In addition, although the final seeds produced under UVB exposure were smaller, their germination rates were unaffected (Day et al., 2001).
In two Antarctic bryophyte species (Sanionia uncinata and Bryum argenteum), no
reductions of net photosynthesis or chlorophyll fluorescence parameters were observed under
current levels of UV-B. However, in the 7 day field UV-B enhancement study, effective
photochemical quantum yield (FPSII) was reduced in S. uncinata. Studies of lichens in
Antarctica have concentrated on the impact of excess visible radiation on photosynthesis and
have found that, while photoinhibition was evident when lichens were water stressed
(Hovenden et al., 1994), it was less likely to be a factor when lichens were fully hydrated
(Kappen et al., 1998a). No significant effects of either screening or supplementation of UV-B
have been observed. As with the excess PAR studies, it appears that lichens are far more
sensitive to moisture content and temperature. Perhaps these factors, which are hard to control
under screening treatments, have tend to obscure any potential impact of UV-B (Huiskes et
al., 2001; Lud et al., 2001b).
113
114
protecting plants against oxidative damage (Bornmann et al., 1997) (Figure 4). Accumulation
in higher plants is primarily in the epidermis, screening physiologically important molecules
below (Robberecht and Caldwell, 1978; Tevini et al., 1991; Vogt et al., 1991; Lois, 1994;
Cuadra and Harborne, 1996; Bjerke et al., 2002; Buffoni-Hall et al., 2002). The accumulation
of UV-B-absorbing pigments would be particularly useful in Antarctic plants because such
passive screens could protect them from UV-B damage when physiological inactivity, due to
desiccation or freezing, renders active repair mechanisms unavailable (Lovelock et al., 1995a,
Lovelock et al., 1995b). UV-absorbing compounds have been investigated in a number of
Antarctic terrestrial species from cyanobacteria to terrestrial plants. In general, cyanobacteria
are protected by mycosporine-like amino acids (MAAs) and scytonemins, while terrestrial
plants contain flavonoids (Rozema et al., 2002). Flavonoids are important UV-B absorbing
pigments, which can be induced within hours in response to UV-B radiation, and are
ubiquitous in higher plants (Cooper- Driver and Bhattacharya, 1998). They have been
extracted from about half of the bryophyte species examined (Markham, 1990). Flavonoids
from herbarium specimens of Antarctic Bryum argenteum were also shown to correlate with
historical ozone levels suggesting the possibility that these were actively induced UV-B
screens (Markham 1990). Recently, high concentrations of UV-B absorbing pigments have
also been reported in two Antarctic mosses, Sanionia uncinata and Andreaea regularis, and
one liverwort, Cephaloziella varians, with positive correlations between pigment
accumulation and flux of natural solar UV-B radiation (Newsham et al., 2002, 2005;
Newsham, 2003). Conversely, UV-B absorbing pigments decreased or showed no change in
response to elevated UV-B levels in seven European, Arctic and South American moss
species (Barsig et al., 1998; Gehrke, 1998, 1999; Searles et al., 1999; Niemi et al., 2002 a, b).
Kaempferol
Quercetin
Figure 4. Molecular structure of kaempferol (above) and quercetin (below) found in some terrestrial and
aquatic higher plants.
115
As a result of these studies, it has been suggested that mosses are less likely to synthesize
UV-B absorbing pigments than other plant groups and are potentially more vulnerable as a
functional type (Gwynn-Jones et al., 1999). Although relatively few mosses have been
studied, negative effects of UV radiation on moss growth and morphology have been reported
for some high latitude species (Sonesson et al., 1996; Searles et al., 1999, 2002; Robson et al.,
2003, Robinson et al., 2005). In many species, flavonoid synthesis is stimulated by exposure
to UV-B radiation (Tevini et al. 1981; Staaij et al. 1995), which seems to act at the gene level
by increasing the expression of the enzymes of the phenylpropanoid pathway (Hahlbrock et
al. 1989; Kubasek et al. 1992) such as Chs, which encodes the enzyme chalcone synthase, and
phenylalanine ammonia lyase (PAL).
Lichens are exposed to the highest irradiance levels when desiccated (Lange et al. 1999),
and in this state, active repair cannot take place. Therefore, lichens are probably more
dependent on UV-B protection by screening compounds than the homeohydric plants. UVprotection in lichens has been tentatively ascribed to secondary compounds, also called lichen
compounds (Fahselt 1994; Rikkinen 1995; Huneck 1999), a heterogeneous group of fungal
compounds with varying biosynthetic pathways (Huneck and Yoshimura 1996). Most lichen
compounds absorb strongly in the UV, and some, such as parietin, additionally absorb
photosynthetically active radiation (PAR) (Solhaug et al. 2003; Hill and Woolhouse 1966).
The widely distributed and common compounds parietin, atranorin, usnic acid, as well as the
structurally less known fungal melanins, are cortical pigments, forming a screen above the
photobiont. Most other compounds are located in the photobiont layer within the upper part of
the medulla (Fahselt and Alstrup 1997). The cortical lichen compounds in particular have a
strong potential to protect the photobiont in lichens against adverse effects of UV-B radiation
(Dietz et al. 2000) found that pigmentation of the cortex, rather than changes in reflectance
and cortex thickness was the most important factor determining percent transmitted PAR in
nine investigated lichen species.
116
Dunn and Robinson 2006; Clarke and Robinson 2008). The fact that these mosses are so well
protected when dry is suggestive of passive protection, as enzymatic repair processes are
unlikely to be active in desiccated organisms (Buffoni-Hall et al. 2003). Passive protection
mechanisms would also be effective when these mosses are frozen and could thus be
particularly beneficial to polar and alpine plants. Protection from UV-B when desiccated
could be due to morphological changes upon drying, which reduce light levels in the cell.
Desiccation tolerant plants typically reduce exposed leaf area when dry by folding or
curling of leaves (Davey and Ellis-Evans 1996; Proctor and Tuba 2002). This reduces
transmission of PAR into the cell by between 40-60% in a range of moss species including B.
pseudotriquetrum and T. ruralis (Seel et al. 1992a). UV-B is likely to be similarly reduced
which would contribute considerable protection at the molecular level. Based on relative turf
densities, desiccation of these three mosses results in reductions in size ranging from 25% in
S. antarctici to 40 to 50% in B. pseudotriquetrum and C. purpureus respectively (Wasley et
al. 2006). When moss cells shrink upon desiccation, cytoplasm volume is reduced,
concentrating cellular contents including UV-B screening compounds and possibly increasing
the attenuation of UV-B. In most ecosystems, periods of high insolation (and associated UVB stress) cause desiccation in bryophytes as they equilibrate leaf turgor with that of their
surroundings (Gehrke 1999). The reverse is true in the Antarctic environment however, where
the major water source is snow-melt, which is maximal during periods of high insolation, and
can coincide with elevated UV-B as a result of ozone depletion. Thus if desiccation is a major
strategy for protection from UV-B these plants may still be at risk from high UV-B during
ozone depletion, especially when this coincides with spring melt.
117
of poly (dA)-(dT) tracts, but not in their rigid centre (Becker and Wang 1989, Lyamichev
1991). CPD formation is less when the bending of the DNA is towards the minor groove
(Pehrson and Cohen, 1992).
One of the transcription factors which have a direct effect on DNA damage formation and
repair is the TATA-box binding protein (TBP). TBP induces the selective formation of 6
4PPs in the TATA-box, where the DNA is bent, but CPDs are formed at the edge of the
TATA-box and outside, where the DNA is not bent (Aboussekhra and Thoma, 1999). Every
CPD acts as a block to transcription and replication, and only a small fraction of dimers
results in a mutation (Britt 1995, 1996).
These DNA lesions, if unrepaired, interfere with DNA transcription and replication and
leading to misreading of the genetic code and leads to mutations and death. Plants uses two
broad strategies to minimise the amount of damage caused by UV-B radiation: decreasing the
penetration of UV-B radiation within the tissue and repairing the damage caused by the
radiation that succeeded in penetrating into the tissue.
One way of decreasing the penetration of UV-B radiation in the tissue is by the
accumulation of UV-absorbing pigments, as described in the previous section. Evidence that
UV-absorbing pigments are important for the protection of the DNA come from various
studies in which plants that are genetically deficient in these pigments were shown to be
much more sensitive to UV-induced DNA damage than their respective wild types (Li et al.
1993; Stapleton et al. 1994).
118
Figure 6. Formation of the UV-induced second most frequently occurring 64 photoproducts and their
Dewar valence isomers. 64 photoproducts are formed at 5_-TC-3_, 5_-CC-3_, 5_-TT-3_ but not at
5_-CT-3_ sites in DNA (cosmobiousa.com).
119
deazariboflavin, with absorption maxima of 380 and 440 nm, respectively) is a lightharvesting antenna absorbing the blue-light photon and transfers excitation energy to the
active (catalytic) cofactor, which is invariably a two electron- reduced flavinadenine
dinucleotide (FADH). Flavin in the excited state donates an electron to the CPD, splitting the
cyclobutane ring, and the electron is transferred back to the flavin concomitantly with the
generation of the two canonical bases (Thoma 1999). CPD photolyases recognize CPDs with
a selectivity similar to that of sequence specific DNA-binding proteins, which suggests that
they could compete with histones for DNA accessibility in a manner similar to transcription
factors (Sancar et. al.1987). Once photolyase binds to CPD, the efficiency of photo
reactivation is extremely high: approximately one dimer split for every blue-light photon
absorbed (Britt 1996).
Although the general light-driven splitting mechanism of photo reactivation is well
understood, a number of aspects of the repair process remain obscure such as (i) how repair
enzymes recognize single DNA lesions with high precision in a structurally heterogeneous
mega base prokaryotic or even chromatin-containing eukaryotic genome, (Roberts 1995,
Reinisch et. al. 1995, Nelson and Bestor 1996) (ii) how the initial reduction of FAD to FADH
takes place.
Figure 7. Formation of the most toxic and mutagenic DNA lesion, cyclobutanepyrimidine dimers by
UV radiation. Dimers can form between two adjacent pyrimidines and their photoreactivation by the
enzyme photolyase in the presence of light.
120
Figure 8. Repair of pyrimidine dimers with photolyase. Energy derived from absorbed light is used to
reverse the photoreaction that caused the lesion. The two chromophores
methenyltetrahydrofolylpolyglutamate (MTHFpolyGlu) and FADH_, perform complementary
functions. On binding of photolyase to a pyrimidine dimer, repair proceeds as follows (Principles of
Lehninger fourth edition).
1 A blue-light photon (300 to 500 nm wavelength) is absorbed by the MTHFpolyGlu, which functions
as a photoantenna. 2 The excitation energy passes to FADH_ in the active site of the enzyme. 3 The
excited flavin (*FADH_) donates an electron to the pyrimidine dimer (shown here in a simplified
representation) to generate an unstable dimer radical. 4 Electronic rearrangement restores the
monomeric pyrimidines, and 5 the electron is transferred back to the flavin radical to regenerate FADH.
Based on site-directed mutagenesis (Heelis et. al.1990, Li et. al.1991) and EPR
investigations (Essenmacher et. al. 1993) it was suggested that the formation of FADH_
results from a temporary photo-reduction and requires an electron transfer from a distant
tryptophan to the light-excited FAD radical quartet state. In addition, by using time-resolved
absorption spectroscopy it has been shown in E. coli DNA photolyase that the excited FAD
radical abstracts an electron from a nearby tryptophan in 30 ps. After subsequent electron
transfer along a chain of three tryptophans, the most remote tryptophan (as a cation radical)
releases a proton to the solvent in about 300 ns, showing that electron transfer occurs before
121
proton dissociation (Aubert et. al 2000). A similar process may take place in photolyase-like
blue-light receptors, (iii) how the enzymes mediate the energy and electron transfer processes
in order to achieve repair with almost maximal efficiency (quantum yield = 0.70.9), and (iv)
in view of the lack of any knowledge of how photolyases recognize their substrate, the
different cleavage rates observed for dimers possessing different configurations and
constitutions remain obscure (Kim et. al 1993).
CONCLUSION
Climate change has already impacted on Antarctic plants. Temperature changes in
Antarctic have led to changes in the distribution of native plants. The current levels of UV-B
have been shown to reduce growth of the plant species, suggesting that ozone depletion may
be having a negative effect on these plants. Although negative effects of UV-B are
ameliorated by UV-screening compounds in many Antarctic plants. Comprehensive
predictions are complicated by both the lack of certainty in the prediction of changes to
abiotic variables, and by the lack of long-term studies investigating recent changes to the
flora. The effects of UV-B radiation on DNA damage and repair in natural environments are
not known clearly as yet and require further research. Furthermore, lichens are usually
regarded as individuals, although they are a symbiotic entity involving at least two partners.
The high resilience of desiccated mosses to DNA damage suggests that passive screening
maybe more important than repair in these species. Further investigations are required on the
effects of UV-B radiation on the DNA of the single symbiotic partners in order to assess
whether the fungus constitutes an effective shield protecting the photobiont against DNA
damage.
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Chapter 6
INTRODUCTION
In Antarctica only cold adapted plants and animals survive, such as Penguins, seals, many
types of lichen, moss and algae. The bryophytes, because of their poikilohydric nature and
alternative strategy of adaption, are one of the very few plant groups which grow in
*
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Antarctica. As such, their role in habitat modification, nutrient cycling, primary production
and providing shelter and security to associated invertebrate animals assume a particular
significance, Incidentally, barring just two species of vascular plants, viz. Deschampsia
antaractica, a grass belonging to family Poaceae, and Colobanthus quitensis, a pearlwort of
family Caryophyllaceae reported to be occurring in Antarctica (Seppelt and Broadly, 1988).
Other groups of plant recorded from the icy continent include lichens, fungi, algae and
bacteria. Thus the Antarctic continent, with its off-lying Islands, is unique in being the only
major landmass almost entirely vegetated by cryptogams, with the lichens predominating in
drier, more exposed situations, while the bryophytes dominant in the more sheltered and
moister habitats. The Antarctic flora in general is impoverished due to both, harsh
environment and isolation of the continent because of vast, cold and turbulent oceanic barrier
of the southern sea. Various biogeographical schemes have been suggested as a means of
classifying Antarctic terrestrial environment but there is currently a general recognition of
three biogeographical zones within the Antarctic as a whole, the sub-, maritime and
continental Antarctic (Smith, 1984; Longton, 1988). The sub-Antarctic zone includes isolated
Islands and archipelagos at high latitudes in the Southern Ocean. Most, with the exception of
South Georgia, Heard and McDonald Islands are close to or north of the oceanic Polar Frontal
Zone. These Islands are under strong maritime influence, which limits and buffers
temperature variation year round, as they are not normally impacted by pack or fast ice.
The Maritime Antarctic is also a region under strong Maritime influence from the
Southern Ocean, in this case with the influence being more seasonal in nature and limited to
the short Antarctic summer period. It includes the western coast of the Antarctic Peninsula to
c. 72 S, the South Shetland, South Orkney and South Sandwich Islands, and the isolated
Bouvetoya and Peter Ioya. The central mountain spine, eastern coast and the more southern
elements of the Antarctic Peninsula are not included, and therefore the term Maritime
Antarctic does not include all or even a majority of the geological region of West Antarctica.
The continental Antarctic is the largest biogeographical zone in terms of area, including
all of East Antarctica, the Balleny Islands, and those parts of the Antarctic Peninsula not
included in the Maritime Antarctic. By contrast with the other two zones, terrestrial habitats
of the continental Antarctic are very limited in extent and more isolated, although they
include coastal rocky regions superficially similar to those of the maritime Antarctic. With
one exception i.e., formation of the extensive ice-free cold deserts of Victoria Land (Lyons et
al., 1997). Air temperature in the continental Antarctic are more extreme than those of the
maritime and sub-Antarctic although, as in all zones, microhabitat temperature may be more
variable, in particular with snow cover giving protection from winter thermal minima. During
the brief summer, absorption of energy by rocks and soil can lead to melting and free water
being available even at the most southern ice-free locations.
The continental Antarctic bryoflora is characterized by just a few species of bryophytes
generally wide distribution belonging to 8-10 genera only, viz. Bryoerythropyllum, Bryum,
Cephaloziella, Ceratodon, Dicranella, Didymodon, Grimmia, Plagiothecium, Pottia and
Sarconeurum. Bryum is the luxirount growing moss genus occurring in Antarctica, because of
great phenotypic plasticity exhibited by it in response to extreme environmental conditions,
coupled with general lack of sporophytes make this as taxonomically most difficult and
confused genus in the continent. Sarconeurum, a monotypic moss genus widely distributed in
Antarctica and Southern South Amrica (Green, 1975; Matteri, 1982), shows the southern
133
most distribution by any bryophyte, being recorded at 82 42'S latitude (Wise and Gressit,
1965).
134
Amblystegiaceae
Andreaeaceae
Bartramiaceae
Brachytheciaceae
Bryaceae
Bryum amblyodon G. Muell- Bipolar, South Shetland Island and Antarctic continent
135
Bryum argenteum Hedw.- Worldwide, South Shetland Island and Antarctic continent
Bryum dichotomum Hedw. Only on Deception Island
Bryum orbiculatifolium Card. Et. Borth Shetland Island
Bryum pallescens Schlieich Ex. Schwaegr- Bipolar, South Shetland Island and
Antarctic continent
Bryum psudotriquetrum (Hedw.) Schwaegr
Bryum urbanskyi Broth (probably= Bryum psudotriquetrum)- South Shetland Island
Pohlia cruda (Hedw) Lindb.- Bipolar, South Shetland Island and Antarctic continent
Pohlia drummondii (C. Muell) A. L. Andrews in Gront [syn: Pohlia inflexa (C.
Muell.) wijk. Et. Marg]- Bipolar, South Shetland Island and Antarctic continent
Pohlia nutans (Hedw) Lindb- Bipolar, South Shetland Island and Antarctic continent
Pohlia wahlenbergii (web. Et Mohr.) Andrews- Bipolar, South Shetland Island
Dicranaceae
Anisothecium cardotii (R. Br. ter.) Ochyra [syn: Dicranella cardotii (R. Brown ter.)
Dixon]- South America, Australia and New Zeland, Subantarctic Island (King
George Island- For a single locality in Ezeurra Inlet. Anisothecium hookeri (C.
Muell.) Broth. [syn: Dicranella hookeri (C.Muell.) Card.]- South Shetland (Only
Deception Island) and Antarctica (Victoria Land) need to revised more carefully
Chorisodontium caiphyllum (Hook. F. et wills.) Broth.- Southern South America,
New Zeland, Antarctic Peninsula and Southern Shetland Island (all the main Island)
Kiaeria pumila (Mitt. In Hook. F.) Ochyra Southern South America, Subantarctic
Island and Southern Shetland Island (only in King George Island), Antarctic
continent
Ditrichaceae
136
Ditrichum lewis and Smithii Ochyra- America, South Shetland (Liringstone and King
George Island) and Antarctic continent.
Encalyptaceae
Grammiaceae
Hypnaceae
137
Meesiaceae
Orthotrichaceae
Plagiotheciaceae
Polytrichaceae
Notoligotrichum trichodon (Hook f. et. Wils.) G.L. Smith- South America and
Antarctica (known only from a single locality on Potter Peninsula, King George
Island)
Polytrichum alpinum (Hedw) G. L. Smith- Bipolar, South Shetland Island and
Antarctic continent
Polytrichum strictum Bird (Syn- P. alpestre Hoppe)- Bipolar, South Shetland Island
and Antarctic continent
Polytrichum juniperinum Hedw- Bipolar, South Shetland Island and Antarctic
continent
Polytrichum piliferum Hedw- Bipolar, South Shetland Island and Antarctic continent
Pottiaceae
138
Syntrichia princeps (De Not.) Mitt. (syn: Tortula princeps De Not)- Bipolar, South
Shetland Island and Antarctic continent
Syntrichia sacicola (Card) zand (syn: Tortulasaricols Card)- Amphiatlantic,
Subantarctic and South Shetland Island
Sarconeurum glaciale (C. Mull.) Card et Bryhm- Tierradel Fuego, Antarctic
continent and South Shetland Island (only in Deception Island)
Sarconeurum tortelloides C.W. Greene
Seligeraceae
139
The aquatic moss vegetation were found at the bottom of lakes from 2.0 to 14.5m in
depth, and were most vigorous from 3 to 5m. In shallow areas, less than 2m in depth, moss
vegetation were not found, probably due to ice action, there were only fragments of algal
mats. Bodin and Nauwerck, (1968), noted that ice movement in summer is the main reason to
explain the absence of aquatic moss colonies in the shallow water zone down to about 2m
depth.
The deepest record of aquatic mosses in the east Antarctica is 32.3m, where P. georgicoantarcticum was found in lake Glubokoye, Schirmacher Oasis, Dronning Maud Land
(Savich-Lyubitskaya and Smirnova, 1964).
140
141
et al., 1992) such as Chs, which encodes the enzyme chalcone synthase, and phenylalanine
ammonia lyase (PAL).
142
Lange and Kappen, 1972). Continental Antarctic bryophytes, whilst less tolerant of increasing
aridity than lichens and also have the ability to survive desiccation. Species-specific
differences in tolerance of desiccation have been detected for three moss species from the
Windmill Islands, East Antarctica (Robinson et al., 2000; Wasley et al., 2006) with two
cosmopolitan species (B. pseudotriquetrum, C. purpureus) able to metabolise at lower turf
water content than the endemic G. antarctici, and S. antarctici. B. pseudotriquetrum also
shows greater plasticity than the other species, with plants from drier sites showing greater
tolerance of desiccation that those from wetter sites, in addition to seasonal changes in
desiccation tolerance (Robinson et al., 2000; Wasley et al., 2006). The ability to survive
repeated desiccation and freezing events is probably related to the high concentrations of
soluble carbohydrates found in these species (Melick and Seppelt, 1994) and in particular the
presence of compounds such as stachyose and trehalose in B. pseudotriquetrum (Robinson et
al., 2000; Wasley et al., 2006). In contrast C. purpureus has a much higher proportion of fatty
acids/soluble carbohydrates. All three species have a high proportion of their fatty acids in
polyunsaturated forms (> 67%), a feature characteristic of cold tolerant higher plants (Zuiga
et al., 1996; Wasley et al., 2006). Quantifying the relative importance of lipids versus soluble
carbohydrates in these freeze tolerant plants stands out as an interesting target for further
study, it may be that lipids are a safer storage compound, since soluble carbohydrates are
known to leak from bryophytes during desiccation-rehydration and freeze thaw cycles
(Melick and Seppelt, 1992).
Water is less likely to be limiting in the relatively moist Maritime Antarctic. On Signy
Island, whilst some xeric species are occasionally water-limited (Davey, 1997c), more
generally photosynthesis is not water-limited (Collins, 1977). When the photosynthetic rates
of a range of xeric and hydric species from this Island were compared under laboratory
conditions, no difference among habitats was detected (Convey, 1994). Maritime moss (A.
regularis, A. depressinervis, P. alpestre, and B. algens) species from a variety of habitats
(hydric, mesic, xeric) also experience increased penetration of light into the turf as drying
occurs, counteracting at least in the short term the loss of productivity during periods of
desiccation (Davey and Rothery, 1996). Water availability has been shown to influence turf
and gametophyte morphology in a range of continental and maritime Antarctic mosses (A.
regularis, P. alpestre, B. algens, Grimmia lawiana and G. antarctici) and this, in turn, can
affect water relations (Nakanishi, 1979). In general, gametophyte shoots are shorter and turf
denser in drier sites (Gimingham and Smith, 1971; Wasley, 2004; Wasley et al., 2006).
Indeed, the changes in plant morphology and growth patterns that are reported as the norm in
many long-term environmental manipulation experiments, often implicitly assumed to relate
primarily to temperature increase, may equally well be explained by changes in microclimate
humidity and soil moisture.
In general, continental Antarctic mosses (G. lawiana, Grimmia plagiopodia, Coscinodon
psudocribrosus and C. bolivianus) can survive repeated freeze-thaw events (Melick and
Seppelt, 1992), whilst maritime species appears to be less tolerant (Davey, 1997b). The
pattern of exposure to freezing is also important - repeated freeze-thaw cycles cause a greater
reduction in gross photosynthesis than constant freezing over the same time period (Kennedy,
1993). Tolerance of freeze-thaw events involves interactions with other environmental
parameters, in particular that of water availability. For example, desiccation before freezing
reduces damage to the photosynthetic apparatus, while protection from freeze-thaw events
can be provided by snow cover acting as an insulator (Lovelock et al., 1995a,b).
143
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Chapter 7
INTRODUCTION
Lichens are one of the most widely distributed groups of organisms in the world; exhibit
their presence in almost all the habitats available. They have an ability to grow on rock,
stones, bark, leaves and various man made substrates including glass plane, iron rods and
plastics. The dry and sterile rock surfaces where other group of plants unable to grow, but
lichens colonizes successfully on them and flourish. Lichens exhibit broadest range of
habitats as they occur both in dry hot and cold desert, from low sea level region to highest
mountains.
*
150
The Antarctic regions is a well known extreme environment for plant life, as the
temperatures are low, there are long periods of frost and snow cover and frequent winds that
causes abrasion and evaporation. Lichens, because of their high resistance to freezing and
their ability to endure long periods of inactivity in a frozen state, can survive in this
environment (Kappen 1973).
In the past few decades a large number of lichenological investigations on Antarctic
lichens were carried out in different regions of the world, thus resulted a fairly clear picture of
the diversity and distribution of lichens in the region. vstedal and Smith (2001) provided the
detailed diversity and biogeography of the lichen biota in Antarctica. Accordingly, South
Georgia which is represented by 194 species of lichen has 47 exclusive species not known
from Antarctic biome. The Antarctic Peninsula has greatest diversity of 268 taxa. The South
Orkeys and South Shethland Islands are represented by 221 and 211 taxa.
Growth
Form
Acarospora. flavocordia Castello and Nimis Cr
Cr
A. gwynnii C.W. Dodge and E.D. Rudolph
Cr
A. macrocyclos Vain.
Cr
A. williamsii Filson
Amandinea coniops (Wahlenb.) M. Choisy ex Cr
Scheid.
- Lecidea coniops Wahenb.
- Buellia coniops (Wahlenb.) Th. Fr.
A. petermannii (Hue) Matzer, H. Mayerhofer Cr
and Scheid.
- Lecanora petermanii Hue
- Rinodina petermanii (Hue) Darb.
- Beltraminia petermannii (Hue) C.W. Dodge
A. punctata (Hoffm.) Coppins and Scheid.
Cr
- Lecidea punctata Hoffm.
- Buellia punctata (Hoffm.) A. Massal.
Distribution
En
En
En
En
Bi
En
Co
10
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12
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20
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22
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Growth
Form
Cr
151
Distribution
Bi
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En
En
En
En
En
En
En
En
En
Bi
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
En
En
En
Co
+
+
+
+
+
+
En
En
Bi
+
+
+
En
En
+
+
En
Bi
En
152
Bi
Cr
Cr
Cr
En
En
Bi
+
+
+
+
+
+
Cr
Cr
Cr
En
En
Bi
+
+
+
Cr
Bi
Cr
Cr
Cr
Cr
En
En
Co
En
+
+
+
+
+
+
Cr
Co
Cr
Cr
En
En
+
+
Cr
Bi
Cr
Co
Cr
Bi
Fo
Co
Fo
Co
Cr
Bi
Fr
Bi
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Bi
Co
+
+
En
Bi
Bi
Bi
Co
+
+
+
+
+
+
Co
En
Co
+
+
+
+
Bi
Co
Co
Bi
+
+
+
-
Co
En
Antarctica
Acarospora badiofusca (Nyl.) Th. Fr.
A. flavocordia Castello and Nimis
A. gwynnii C.W. Dodge and E.D. Rudolph
A. macrocyclos Vain.
A. williamsii Filson
India
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7
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Antarctica
Amandinea coniops (Wahlenb.) M. Choisy ex Scheid.
A. petermannii (Hue) Matzer, H. Mayerhofer and Scheid.
A. punctata (Hoffm.) Coppins and Scheid.
Arthorhaphis alpina (Schaer.) R. Sant.
Arthonia molendoi (Frauenf.) R. Sant.
A. rufidula (Hue) D. Hawksw., R. Sant. and vstedal
Bacidia johnstonii C.W. Dodge
B. stipata I.M. Lamb
Bryonora castanea (Hepp) Poelt
Buellia darbishirei I.M. Lamb
B. frigida Darb.
B. grimmiae Filson
B. grisea C.W. Dodge and G.E. Baker
B. illaetabilis I.M. Lamb.
B. lingonoides R. Filson
B. pallida Dodge and Baker
B. papillata (Sommerf.) Tuck.
B. pycnogonoides C.W. Dodge and G.E. Baker
B. subfrigida May. Inoue
Caloplaca athallina Darb.
C. cerina (Ehrh. ex Hedw.) Th. Fr.
C. citrina (Hoffm.) Th. Fr.
C. exsecuta (Nyl.) Dalla Torre
C. holocarpa (Hoffm.) Wade
C. frigida Schting
C. lewis-smithii Schting and vstedal
C. saxicola (Hoffm.) Nordin
Candelaria murrayi Poelt
Candelariella aurella (Hoffm.) Zahlbr.
C. flava (C.W. Dodge and G.E. Baker) Castello and Nimis
C. vitellina (Ehrh.) Mll. Arg.
Carbonea assentiens (Nyl.) Hertel
C. vorticosa (Flrke) Hertel
Cetraria aculeata (Schreb.) Fr.
C. islandica (L.) Ach.
Chrysothrix chlorina (Ach.) J.R. Laundon
Cladia aggregate (Sw.) Nyl.
Cladonia bellidiflora (Ach.) Schaer
C. carneola (Fr.) Fr.
C. chlorophaea (Flrke ex Sommerf.) Spreng.
C. deformis (L.) Hoffm.
C. fimbriata (L.) Fr.
C. galindezii vstedal
India
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+
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+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
-
Antarctica
C. gracilis (L.) Wild.
C. mitis Sandst.
C. phyllophora Hoffm.
C. pleurota (Flrke) Schaer.
C. pocillum (Ach.) O.J. Rich.
C. pyxidata (L.) Hoffm.
C. rangiferina (L.) Webber
C. scabriuscula (Delise) Nyl.
C. squamosa (Scop.) Hoffm.
C. subsubulata Nyl.
Collema tenax (Sw.) Nyl.
Lecania racovitzae (Vain.) Darb.
Lecanora epibryon (Ach.) Ach.
L. expectans Darb.
L. frustulosa (Dicks.) Ach.
L. fuscobrunnea C.W. Dodge and G.E. Baker
L. geophila (Th. Fr.) Poelt
L. mawsonii C.W. Dodge
L. mons-nivis Darb.
L. orosthea (Ach.) Ach.
L. polytropa (Hoffm.) Rabenh.
L. sverdrupiana vstedal
Lecidea andersonii Filson
L. atrobrunnea (Ramond) Schaer.
L. auriculata Th. Fr.
L. cancriformis C.W. Doge and G.E. Baker
L. lapicida (Ach.) Ach.
L. placodiiformis Hue
Lecidella elaeochroma (Ach.) M. Choisy
L. siplei (C.W. Dodge and G.E. Baker) May Inoue
L. stigmatea (Ach.) Hertel and Leuckret
Lecidoma demissum (Rutstr.) Gotth. Schneid. and Hertel
Lepraria cacuminum (A. Massal.) Kmmerl. and Leuckert
L. neglecta (Nyl.) Erichs.
Leproloma vouaux (Hue) J.R. Laundon
Megaspora verrucosa (Ach.) Hafellner and V. Writh
Ochrolechia tartarea (L.) A. Massal.
Parmelia saxatilis (L.) Ach.
P. sulcata Taylor
Peltigera didactyla (With.) J.R. Laundon
P. rufescens (Weis) Humb.
Pertusaria coccodes (Ach.) Nyl.
Phaeophyscia endococcina (Krb.) Moberg
Phaeorrhiza nimbosa (Fr.) H. Mayrhofer and Poelt
Physcia caesia (Hoffm.) Furner.
155
India
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95
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115
116
117
118
119
120
121
122
123
124
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Antarctica
P. dubia (Hoffm.) Lettau
Physconia muscigena (Ach.) Poelt
Placynthiella icamalea (Ach.) Coppins and P. James
Pleopsidium chlorophanum (Wahlenb.) Zopf.
Protoparmelia badia (Hoffm.) Hafellner
Pseudephebe minuscula (Nyl. Ex Arnold) Brodo and Hawks.
Psilolechia lucida (Ach.) M. Choisy
Rhizocarpon badioatrum (Flrke ex Spreng.) Th. Fr.
R. disporum (Hepp) Mll. Arg.
R. geminatum Krb.
R. geograhicium (L.) DC
R. nidificum (Hue) Darb.
R. superficiale (Schaer.) Malme
Rhizoplaca melanophthalma (Ram.) Leuckert and Poelt
Rinodina endophragmia I.M. Lamb.
R. olivaceobrunnea C.W. Dodge and G.E. Baker
Sarcogyne privigna (Ach.) A. Massal
Sporastatia testudinea (Ach.) A. Massal.
Stereocaulon alpinum Laurer
Tephromela atra (Huds.) Hafellner ex Kalb
Umbilicaria africana (Jatta) Krog and Swinscow
U. antarctica Frey and I.M. Lamb
U. aprina Nyl.
U. decussata (Vill.) Zahlbr.
U. krascheninnikovii (Savicz) Zahlbr.
U. thamnodes Hue
Usnea antarctica Du. Rietz
U. sphacelata R. Br.
Verrucaria maura Wahlenb. in Ach.
V. muralis Ach.
Xanthoria candelaria (L.) Th. Fr.
X. elegans (Link) Th. Fr.
X. mawsonii Dodge
India
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+
+
+
+
+
+
157
South Shetland, South Georgia and outlying of Falkland Islands. Some members of
the stipitate and crustose members of lichens also exhibit their restricted distribution
in the west coast of the Antarctic Peninsula, South Shetland and South Orkney
Islands are Bacidia stipitata, Catilaria corymbosa, Caloplaca regalis, Lecania
brialmontii and Rinodina petermanii. The Antarctic Peninsula (west coast) exhibit
occurrence of distinctive elements such as Cladonia rangiferina, Cornicularia
aculenta, Cystocoleus niger, Himantormia lugubris, Massalengia cornosa, Pannaria
hookeri, Parmelia gerlachei, P. ushuaiensis, Pseudophebe pubescens,
Sphaerophorus globosus and Stereocaulon glabrum. Most of the species under this
category exhibit a strong maritime tendency.
2. The Antarctic Peninsula (east coast): The Antarctic Peninsula possesses a continuous
mountainous range of altitudinal variation of 1500 2500 m in height. Due to the
height of the mountain range the climate divide between two coasts. The west coast
has a strong maritime climate with relatively warm, wet winds but the east coast
having extensive ice shelf provide a relatively cold, dry winds approaching that of
continental Antarctica. Usnea sulphurea and U. fasciata dominates the area.
Pseudophebe minuscule also occur in this region.
3. The Antarctic Peninsula Endemic: The stipitate crustose lichen species such as
Bacidia stipata, Catilaria corymbosa, Caloplaca regalis, Lecania brialmontii and
Rinodina petermanii are restricted to the west coast of the Antarctic Peninsula and
South Shetland and South Orkeney Islands and have originated in this region
(Lindsay 1975).
4. The Circumpolar Antarctic: Buellia frigida an endemic species of Antarctica and
Xanthoria elegans a circumpolar species exhibit their scattered occurrence in this
category. Though the Arctic and Antarctic lichens exhibit many similarities in their
origins, however, the restricted nature of Antarctic lichen flora show little similarity
with the Arctic. Out of the 45 genera of lichens known from the Arctic 20 are also
known from the Antarctic. Out of 328 species of lichens recorded from Greenland
(Lynge 1937), 47 are found in the Antarctic, while out of the 65 genera listed from
the former region 46 are found in the later. The greater number of genera in common
between Arctic and Antarctic supports the idea that at one time the polar lichen floras
were basically similar, but the geographical isolation, especially that of Antarctica
from other southern hemisphere continents, has led to differences through speciation.
158
Figure 1. Map showing the location of Schirmacher Oasis and surrounding nunataks.
The National Botanical Research Institute (NBRI-CSIR) Lucknow, was identified by the
Ministry of Earth Science, New Delhi as the centre for carrying out lichenological studies in
SO in the year 1991. The organization participated for the first time in the 11th Indian
Antarctic Expedition (IAE) during the year 1991-92 and reported a total of 26 lichen species
(Upreti and Pant 1995, Upreti 1996, 1997). During 17th IAE Pandey and Upreti (2000) listed
a total of 19 lichen species collected from the SO and the Vettiya nunatak. Mean while Gupta
et al. (1999) of the Botanical Survey of India also collected some lichens form SO during 18th
IAE and reported five more species. NBRI continued its participation in the 22nd IAE and
more extensively and intensively explored the 31 sites of the whole stretch of SO together
with seven nunataks located nearby and listed a total of 35 lichen species (Nayaka and Upreti
2005).
The consolidation of published accounts on the SO lichens revealed the occurrence of
total of 48 species in the area (Nayaka et al. 2009). Out of the 48 lichen species that occur in
SO, 19 are cosmopolitan in distribution. Olech and Singh (2010) during 23rd IAE in 2003-04
made a detailed lichenological survey of the Oasis and reported 57 lichen taxa after adding 22
new addition to the Oasis. The consolidation of all the available lichenological studies
revealed the occurrence of 69 species in the SO (Table 1). The species of lichen genus Buellia
dominates the area with 10 species, followed by 9 species of Lecanora, 5 species each of
Caloplaca and Umbilicaria. The crust forming species dominates the area with 54 species,
while only 9 species are foliose, 4 are fruticose and single species of leprose form.
159
the affinities of the lichen flora of the subcontinent with neighbouring area and other regions
including Antarctica. A large number of lichen species of the subcontinent are cosmopolitan
in distribution. The eastern Himalayan lichen flora has many species common with the SinoJapanese and south-east Asian regions. Lichen species present in the western Himalayas
exhibit a distinct affinity with the lichen flora of northern Europe. Though the Antarctic
lichen flora exhibits a more restricted nature, however most of the lichen genera are known
from Antarctica is also exhibit their occurrence in other regions of the world including Indian
subcontinent. The SO is represented by the following 23 genera of lichens; Acarospora,
Amandinea, Arthonia, Bacidia, Buellia, Caloplaca, Candelaria, Carbonea, Lecania,
Lecanora, Lecidea, Lecidella, Lepraria, Physcia, Pleopsidium, Pseudophebe, Rhizocarpon,
Rahizoplaca, Rinodina, Sarcogyna, Umbilicaria, Usnea and Xanthoria. Except the genus
Pseudophebe all the 22 genera know from SO are also know from India and the Indian
subcontinent. Out of the 69 species, so far known from SO, 19 species belonging to 13 genera
are also known from India. The lichen taxa common between India and SO are Amandinea
punctata, Caloplaca cerina, C. citrina, C. frigida, C. saxicola, Carbonea vorticosa, Lecanora
polytropa, Lecidea auriculata, L. lapicida. Lecidella stigmatea, Physcia caesia, P. dubia,
Pleopsidium chlorphanum, Rhizocarpon geographicum, Rhizoplaca melanophthalma,
Sarcogyne privigna, Umbilicaria decussata, U. vellea and Xanthoria elegans. In India most
of the lichen species which exhibit their occurrence in SO are mostly exhibit their restricted
distribution in temperate and alpine regions of the Himalayas. Rhizocaron geographicum
sometimes also found growing on rocks in higher altitudes of southern Indian region. Out of
439 taxa of lichens so far known from Antarctica and South Georgia (vstedal and Smith
2001, 2004) 76 species exhibit similarities with Indian subcontinent and 68 with the SO. Out
of 76 Indian subcontinent lichen species common to Antarctica (Figure 2) the lichen genus
Cladonia is represented by 64 species in the subcontinent, out of which 17 (47%) species are
also known to occur in Antarctica. However, the SO is devoid of this genus as not a single
species of Cladonia is known from there.
Lecidea
5%
Lecidella
5%
Parmelia
5%
Physcia
5%
Xanthoria
5%
Caloplaca
12%
Cladonia
47%
Rhizocarpon
8%
Umbilicaria
8%
160
The species of Phaeophyscia, Physcia and Physconia are widely distributed in Indian
Himalayan regions are also known from Antarctica, while except for Physcia other two
genera do not exhibit their representation in SO. Most of Indian species of Rhizocarpon show
affinities to the Antarctic lichens as 5 species are commonly known from both the regions.
It is interesting to note that Buellia, a crustose thalloid genus, most common in Antarctica
and Indian subcontinent have different species, but not a single species of Antarctica occur in
India. However, SO is represented 10 species of Buellia known from Antarctica. Most of the
genera of lichens known from India are also reported from Antarctica. The greater number of
genera is common between Indian subcontinent and Antarctica supports the idea that one time
the Indian subcontinent and Antarctic lichen flora were basically similar, but the geographical
isolation that of Antarctica from other southern hemisphere continents has led to difference
through speciation.
ACKNOWLEDGMENTS
We thank Director, National Botanical Research Institute, Lucknow for providing
laboratory facilities, Ministry of Earth Science and Council of Scientific and Industrial
Research, New Delhi for facilitating Antarctic expeditions.
REFERENCES
Awasthi, D.D.: Lichenology in Indian Subcontinent. Bishen Singh Mahendra Pal Singh,
Dehra Dun (2000).
Golubkova, N.S. and I.M. Simonov: Lishayniki Oazisa Shirmakhera. Trudy Sovet Skoy East
Antarkti Cheskoy Eksfeditsii Leningrad, 60, 317-327 (1972).
Gupta, B.K., G.P. Sinha, and D.K. Singh: A note on lichens of Schirmacher Oasis, East
Antarctica. Indian J. Forestry., 22(3), 292-294 (1999).
Kappen, L.: Response to extreme environments. In: The lichens (Eds: V. Ahmadjian and
M.E. Hale). Academic Press, New York. pp 311-380 (1973).
Lindsay, D.C.: Lichens of cold deserts. In: Lichen Ecology (Eds: M.R.D. Seaward.).
Academic Press, London. pp. 183-209 (1977).
Lindsay, D.C.: The macrolichens of South Georgia. Brit. Antarct. Surv. Sci. Rep., 89, 1-91
(1975).
Lynge, B.: Lichens from West Greenland, collected chiefly by Th. M. Fries. Meddr.
Grnland., 118, 1-225 (1937).
Nayaka, S. and D.K. Upreti: Schirmacher Oasis, East Antarctic, a lichenologically interesting
region. Curr. Sci., 89(7), 1059-1060 (2005).
Nayaka, S., D.K. Upreti and R. Bajpai: Diversity and adaptive response of lichens in
Antarctica with special reference to Schirmacher Oasis. In: Frontiers in Fungal Ecology,
Diversity and Metabolites (Ed. K.R. Sridhar.). I.K. Internataional Publishing House Pvt.
Ltd., New Delhi. pp. 107-123 (2009).
Olech, M. and S.M. Singh: Lichens and Lichenicolous Fungi of Schirmacher Oasis,
Antarctica. National Centre for Antarctic and Ocean Research, Vasco da Gama (2010).
161
vstedal, D.O. and R.I.L. Smith: Addition and corrections to the lichens of Antarctica and
South Georgia. Cryptogamie Mycologie. 25(4), 323-331 (2004).
vstedal, D.O. and R.I.L. Smith: Lichens of Antarctica and South Georgia. A guide to their
identification and ecology. Cambridge University Press, U.K. (2001).
Pandey, V. and D.K. Upreti: Lichen flora of Schirmacher Oasis and Vettiyya Nunatak. In:
Scientific Report: Seventeenth Indian Expedition to Antarctica. Ministry of Earth
Science, New Delhi. Technical Publication No. 15. pp. 185-201 (2000).
Ritcher, W.: Biology. In: The Schirmacher Oasis, Queen Maud Land, East Antarctica and its
surroundings (Eds: P. Bormann and D. Fritzsche.). Germany. 321-347 (1995).
Ritcher, W.: The lichens of the Schirmacher Oasis (East Antarctica). Geodtische und
Geophysikalische Verffentlichungen, Reihe 1, Berlin 16, 471-488 (1990).
Rudolph, E.D.: Lichen distribution. In: Terrestrial Life in Antarctica (Eds: V. Bushnell).
Antarct. Map Fol. Ser., 5, 9-11 (1967).
Upreti, D.K. and G. Pant: Lichen flora in and around Maitri region, Schirmacher Oasis, East
Antarctica. In: Scientific Report: Eleventh Indian Expedition to Antarctica. Ministry of
Earth Science, New Delhi. Technical Publication No. 9, pp. 229-241 (1995).
Upreti, D.K.: Lecideoid lichens from the Schirmacher Oasis, East Antarctica. Willdenowia.,
25, 681-686 (1996).
Upreti, D.K.: Notes on some crustose lichens from Schirmacher Oasis, East Antarctica.
Feddes Repertorium., 108(3-4), 281-286 (1997).
Chapter 8
164
INTRODUCTION
Lichen is a composite plant consisting of symbiotically associated fungal (mycobiont)
and algal or cyanobacterial (photobiont) partners. The mycobiont is usually an ascomycete
and in very few cases a basidiomycete, and a photobiont is usually a green alga but in about
10% of lichens a cyanobacterium. Lichens are strictly dependent on ambient moisture for
their metabolic activities. Unlike homiohydric vascular plants lichens lack stomata, cuticle
and water transport system such as roots, xylem vessel and tracheids. Hence, they can not
actively regulate their water content and are called as poikilohydrous organism. The
homiohydric organisms are desiccation sensitive and die when water content falls below a
certain threshold, where as poikilohydric organism include both desiccation-sensitive and
desiccation-tolerant species. Desiccation-tolerance is the ability to revive from the air-dried
state when water is provided. It is also found in prokaryotes, algae and bryophytes, and
occasionally in pteridophytes, but very rare in the vegetative tissues of angiosperms or in
animal tissues. The vast majority of lichens are desiccation-tolerant. Under natural conditions,
the life of most lichens is exposed to rapidly changing water contents and correspondingly
rapidly changing physiological activity such as respiration and photosynthesis (Kranner et al.
2008). Antarctica is a land of extremes. About 98% of the continent is permanently covered
with ice, while remaining 2% of ice-free land is restricted mostly to the periphery of the
continent. These ice-free areas are the only habitat available for the growth lichens.
Antarctica is the coldest continent with heat balances is everywhere negative (Engelskjn
1986). It is the windiest of the continents with the highest wind speed measured at d'Urville
(327 km/h). The wind can act in three ways, by desiccation, wind-chill and ice-blast. The
absolute humidity in the continent is 0.03%, lower than that of the Sahara Desert. Low
average humidity combined with the extreme cold make the South Pole region the world's
driest desert. It rains very rarely in maritime Antarctica, otherwise the snowfall, which
averages less than one inch annually is the major precipitation in the continent. The sun does
not shine at the South Pole for six months of the year. When the sun does shine, much less
solar energy actually reaches the ground at the Pole because the sun's rays pass through a
thicker layer of atmosphere than at the Equator. Also, due to the predominance of ice and
snow covering Antarctica, most of the sun's rays that do reach the ground are reflected back
into space leaving the continent cold with un-molten ice mass. The freshwater lakes in
Antarctica are mostly confined to certain ice-free areas as Larsemann Hills, Schirmacher
Oasis, Bunger Hills and Vestfold Hills. These lakes, which remain frozen for most of the
year, are fed by glacier and snowmelt streams during the short austral summer months.
Despite the restricted availability of liquid water and other extremes a total of 439 lichens do
exist in Antarctica (vstedal and Smith 2001; 2004), which can be directly attributed to their
adaptation to the stressor (Kappen 2000). The adaptive stimulus is clearly expressed in
lichens as they are the most successful elements of the terrestrial biota in Antarctica and have
obvious dominance in ice-free areas. They are superior to bryophytes and perhaps also to free
living algae and fungi, particularly in harsher regions (Kappen 2004). Lichens together with
bryophytes or alone form the largest amount of standing biomass in Antarctic landscapes with
up to 950 kgm-2 in continental and up to 1300 gm-2 in maritime Antarctic habitats (Kappen
2000). Nayaka et al. (2009) discussed in detail the adaptive response of Antarctic lichens to
various stress in factors. The water being a major constrain in Antarctica physiological
165
adaptation to it is an interesting aspect for research. In the present study water relations of six
common Antarctic lichens growing in different water regime in Schirmacher Oasis (SO) were
studied in details. The aim of the study is to identify a lichen species that is physiologically
better adapted to water stress in Antarctica. The aim is achieved by through psychrometric
technique and measuring water holding capacity. This technique relies on water potential data
from which pressure volume (PV) curve is constructed and various isotherms are derived. The
PV isotherm is one of the widely used tools utilized for characterizing water status (Becket
1997). The PV isotherm has proved useful in assessing desiccation tolerance and the
distribution of tree species in Malay-Thai peninsula (Baltzer et al. 2008). Many useful
relations such as osmotic potential (OP) at full turgor, turgor loss point (TLP), relative water
content (RWC) at TLP, elasticity modulus, solute concentration, symplastic, apoplastic and
intercellular water contents can be measured with the help of PV curve.
Table 1. Antarctic lichen taxa analyzed for the water relation and their ecology
1
2
3
4
5
6
Sample
Habit
Habitat
Substratum
Crustose
Crustose
Foliose
Water drainage
Dry area
Dry area
Rock
Moss
Soil
Exposure
to sun
Exposed
Partial shade
Exposed
Foliose
Foliose
Foliose
Water drainage
Dry area
Dry area
Rock
Rock
Rock
Exposed
Exposed
Partial shade
Locality
Priyadarshini Lake
Near Circle lake
North West of
Maitri
Priyadarshini Lake
Vetehia nunatak
Trishul Hill
Table 2. Water holding capacity and derivatives of PV curve for Antarctic lichens
Sample
1
2
3
4
5
6
B. frigida
L. epibryon
R. melanophthalma
U. aprina
U. decussata
X. elegans
Water holding
capacity (% dry wt.)
70.9 110.57
90.62 145.24
120.83 187.2
130.6 229.12
115.51 220.62
148.42 208.22
OP at full Turgor
(MPa)
-1.02 0.46
-0.90 0.26
-0.90 0.29
-0.64 0.25
-1.63 0.22
-0.85 0.80
Apoplastic
water fraction
12.0 5.66
19.75 7.50
12.29 7.34
28.0 8.3
21.38 9.24
21.0 5.31
Elasticity
modulus (MPa)
6.42 0.43
15.01 5.12
3.98 2.95
5.75 3.03
4.55 2.59
9.50 4.50
Water content
at TLP
61.75 1.5
48.25 5.91
67.43 8.02
66.25 3.4
58.75 8.46
56.25 2.2
167
is found growing luxuriantly on rocks of Vitteheia and Baalsrudfjellet nunatak along with
Pseudophebe minuscuala (Nyl. ex. Arnold) Brodo and D. Hawksw. Xanthoria elegans (Link)
Th. Fr. It is a foliose lichen, up to 5 cm in diam., orange in colour with radiating lobes. It is
a cosmopolitan species very common in Antarctica. It is an attractive and very prominent due
to its bright orange colouration. But in SO it is rare and found on rocks, in partially shaded,
shelf facing side of Trishul Hill.
168
The PV curve reconstructed using RWCc and the OP () at full turgor is noted as the yintercept of the linear portion of the PV curve (Figure 1). Regression line going through this
linear portion of the curve intercepts at x-axis and yields the symplastic and apoplastic
fraction of water. Tissue elasticity was calculated from the relationship between and RWCc
(Stadelmann 1984). The bulk elasticity modulus of tissue expresses the change in turgor of
tissue cells for a unit change in the relative water content of the cells (e = dP/dr).
RESULTS
The water holding capacity of Antarctic lichen varied from 70.9 229.12 % of dry of wt.
(Table 2). The foliose lichen U. aprina, which has larger, thick thallus and grows on rock in
lake drainages had maximum water holding capacity ranging from 130.6 229.12 % dry wt.
However, a crustose lichen B. frigida, found in the same habitat but having smaller thallus has
lower water holding potential which ranged from 70.9 110.57 % of dry wt. In general
foliose lichens exhibited better water holding capacity in comparison to crustose.
169
DISCUSSION
The lichens, being exposed to frequent water stress have developed high degree of water
holding capacity. They lack cuticle to impede entry of water in to the thallus. The spongy
nature of the thallus tissue, presence of large intercellular spaces, the gelatinous sheath of
algal cells and swollen hyphal wall allows them to hold good amount of water. The thalli
often can absorb considerably more water during a prolonged immersion of several hours.
The lichens are hygroscopic in nature and absorb moisture from the surroundings even at low
humidity. The lichen can be hydrated and activated quickly by the snow, dew, mist or glacier
meltwater and resume photosynthesis. Some lichens such as Umbilicaria having dark
coloured thalii by absorbing sunlight become much warmer and can melt snow crystals
deposited over them. Further, moistened thalli can reach an over-air temperature of nearly 23
K, which means that they can become as warm as 20C at maximum that induce maximal
productivity. Some lichens are capable of gaining enough moisture from the humidity at the
edge of a temporary snow patch on the rock without any visible melting process. Blum (1973)
enumerated the water content (water holding capacity) of saturated thali of some lichens
which ranged from 116 738 % of dry wt. (maximum in cyanolichen Collema flaccidum
(Ach.) Ach.). Umbilicaria pustulata (L.) Hoffm. held 201 % of dry wt. of water, in 5 min. of
immersion, 203 % in 1 hr and 209 % in 15 hr. Similarly, Xanthoria parietina (L.) Beltr. held
178, 188 and 227 % of dry wt. of water in 5 min, 1 and 15 hr respectively. However, in case
of Antarctic U. aprina, U. decussata and X. elegans their thallus reached full saturation within
30 min. of immersion with maximum of 229.12, 220.62 and 208 % of dry wt. of water
respectively. This indicates the quicker response of Antarctic lichens when water is made
available and an adaptation to combat water scarcity. The water holding capacity of crustose
lichens in both the regions, i.e. main land (Blum 1973) as well as in Antarctica was lesser
compared to foliose lichens.
Unlike vascular plants, lichens can stay longer in desiccation state and their metabolic
activity can recover after their tissues have been reduced to very low RWC. Such plants are
called resurrection plants and have evolved desiccation tolerance (Alamillo and Bartels
2001, Bartels 2005; Tuba et al. 1998). The lichen can photosynthesize at low water potentials
such as 20 MPa at 20C. Experiments in hot deserts have shown that lichens are able to
photosynthesize even at low water potentials as low as 38 MPa (Nash et al. 1990). As soon
as availability of moisture stops lichens thallus quickly desiccates, loses up to 97% of its
water, and falls into an anabiotic state. Under low water potential photosynthesis rate will be
low and can be considered as a means of keeping the photosynthetic apparatus intact and
170
producing frost-protective carbohydrates than for actual dry matter. The psychrometric
technique explores new possibilities to elucidating the water status of plants including
cryptogams such as lichens, bryophytes, pteridophytes. According to several studies utilizing
PV curve, an ideal desiccation tolerant lichen would have low OP at full turgor, lesser
apoplastic water content, low RWC at turgor loss and more importantly less elasticity
modulus. The low OP at full turgor indicates that the cell sap has low salt concentration and
water potential would be high or near to zero. The lower apoplastic contents mean little
quantity of water is present in the cell wall pores and more within in the cell (symplast),
which contributes to high water potential. It is always necessary for a desiccation tolerant
plant that it maintains turgidity of the cell even at lower amount of water. Hence, low RWC at
turgor loss would be beneficial. Similarly, elasticity modulus indicates the stretchability of the
cell wall. The low elasticity modulus value means high elasticity of cell wall and more
tolerance for desiccation. Becket (1997) studied the PV isotherm of several poikilohydric
plants including lichen Roccella hypomecha (Ach.) Bory., and also provided explanation for
varied values of PV isotherms. The lichen R. hypomecha consisted low OP, low RWC at
turgor loss, low elasticity modulus and low solute concentration. Proctor et al. (1998) studied
the water contents of several bryophytes including a lichen Cladonia convoluta (Lam.) Cout.,
using PV relationship, where the lichen contained low OP, low RWC at TLC and low bulk
elasticity modulus. In the present study all the lichens studied exhibited varied values for
measured parameters. U. decussata had lowest OP (1.63 0.22 MPa) at full turgor, but
slightly higher water content at TLP (58.75 8.46) and apoplastic water (21.38 9.24). B.
frigida had lowest apoplastic water fraction (12.0 5.66), but higher OP at full turgor (-1.02
0.46 MPa). Similarly, L. epibryon had lowest water content at TLP (48.25 5.91), but
higher elasticity modulus (15.01 5.12 MPa), apoplastic water content (19.75 7.50) and OP
at full turgor (-0.90 0.26). Hence, it is has become necessary to relay on one strong
parameter, compare other parameter accordingly and decide the desiccation tolerance of the
lichens. The elasticity modulus is thought to be determined by the mechanical properties of
cell walls (Cheung et al. 1975, Tyree and Hammel 1972). In leaf cells that have turgor
elasticity modulus has a critical role in water relations. Advantage of the small elasticity
modulus in the drought environment is that it contributes to the turgor maintenance of the leaf
cells under conditions of low water content (Kozlowski et al. 1991). Hence, in the present
study desiccation tolerances of lichens are compared based on their elasticity modulus. The
lichens growing on moraine or rock in exposed areas are better desiccation tolerant and have
lesser elasticity modulus. Among them R. melanophthalma exhibited lowest (3.98 2.95)
elasticity modulus indicating its highest desiccation tolerance. It is followed by U. decussata
(4.55 2.59), U. aprina (5.75 3.03) and B. frigida (6.42 0.43). It is obvious that lichens
growing in exposed areas are the one who first experience the extremes of Antarctic
conditions. They lose water either due to high irradiation, wind or low temperature and such
conditions are very frequent in Antarctica. Hence, the cells of the lichens probably have
become more stretchable or elastic. Where as in X. elegans, a lichen growing in partial shade
at Trishul hill elasticity modulus is moderate (9.50 4.50) indicating its medium level of
desiccation tolerance. L. epibryon, a muscicolous lichen that grows in partial shade has
highest elasticity modulus. The moss cushions acts like sponge and hold water for longer
duration. Further, the mosses have many dead cells in their leaf tissue which again holds
water. The moss in Antarctica usually grows in shaded area, between or under the rocks. The
lichen by growing over them can obtain water for longer duration and hence developed less
171
desiccation tolerance. In general, by considering all water relation parameters studied the
foliose lichens R. melanophthalma can be considered as highly desiccation tolerant species.
As discussed earlier it has low elasticity modulus, but also has comparatively lower low OP at
full turgor (0.90 0.29 MPa), lower apoplastic water fraction (12.29 7.34) and lower
water content at TLP (67.43 8.02). Further, water holding capacity of R. melanophthalma is
considerably high which ranges from 120.83 187.2 % of dry wt.
CONCLUSION
Among all the theories, the adaptation theory explains best the dominance of lichens in
Antarctica. The lichens are adapted both morphologically as well as physiologically to the
stress components of the continent. The water relation study carried out with the help of
psychrometric method clearly indicates the differential levels of desiccation tolerance in
Antarctic lichens. The habitat selection and restricted distribution of lichens is a clear strategy
for the survival. Hence, X. elegans is found only on shelf facing side of Trishul hill in SO
which receives shade at least for half a day during summer. The muscicolous lichen L.
epibryon grows only on moss so that it gets water for longer duration. As per elasticity
modulus desiccation tolerance is higher in sun exposed lichens, moderate in partially shaded
and least in muscicolous. Hence, desiccation tolerance in Antarctic lichens is of following
sequence; R. melanophthalma > U. decussata > U. aprina > B. frigida > X. elegans > L.
epibryon. Further, water absorbance ability is faster and water holding capacity is higher
which reaches up to 200% of their dry weight. Apparently, lichens did not develop unique
adaptive mechanisms in the extreme Antarctic environment. The high desiccation tolerance
and some morphological features of Antarctic lichens are also been observed in lichens
growing in harsh climate of elsewhere (Kappen, 1988). However, the ability to tolerate stress,
survive and evolve is high in Antarctic lichens.
ACKNOWLEDGMENTS
We are thank full to the Director, National Botanical Research Institute for providing the
necessary facilities, to Dr. U.V. Pathre for allowing us to utilize facilities of Plant Physiology
Laboratory, to Council of Scientific and Industrial Research, New Delhi for financial
assistance, to National Centre for Antarctic and Ocean Research, Vasco da Gama for
selecting S.N. for 28th Indian Antarctic Expedition, to the leaders and members of the
expedition for their cooperation during the collection of lichen samples, to Dr. Ajit Pratap
Singh and members of Plant Physiology Laboratory for their cooperation during the study.
REFERENCES
Alamillo, J. and D. Bartels: Effects of desiccation on photosynthesis pigments and the ELIPlike dsp 22 protein complexes in the resurrection plant Craterostigma plantagineum.
Plant. Sci., 160, 1161-1170 (2001).
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Baltzer, J.L., S.J. Davies, S. Bunyavejchewin and N.S.M. Noor: The role of desiccation
tolerance in determining tree species distributions along the Malay-Thai Peninsula.
Funct. Ecol., 22, 221-231. (2008).
Bartels, D.: Desiccation tolerance studied in the resurrection plant Craterostigma
plantagineum. Integrative Comparative Biology., 45, 696-701 (2005).
Beckett, R.P.: Pressure volume analysis of a range of poikilohydric plants implies the
existence of negative turgor in vegetative cells. Annals of Bot. 79, 145-152 (1997).
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(1995).
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Cheung, Y.N.S., T.M. Tyree and J. Dainty: Water relations parameters on single leaves
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(2000).
Kappen, L.: The diversity of lichens in Antarctica, a review and comments. Biblioth.
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Kozlowski, T.T., P.J. Kramer and S.G. Pallardy: The physiological ecology of woody plants.
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Kranner, I., R.P. Beckett, A. Hochman and T.H. Nash III.: Desiccation tolerance in lichens: a
review. Bryol. 111(4), 576-593 (2008).
Nash III, T.H., A. Reiner, B. Demmig-Adams, E. Kilian, W.M. Kaiser and O.L. Lance: The
effect of atmospheric desiccation and osmotic water stress on photosynthesis and dark
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Nayaka, S., D.K. Upreti and R. Bajpai: Diversity and adaptive response of lichens in
Antarctica with special reference to Schirmacher Oasis. In: Frontiers in Fungal Ecology,
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Ltd., New Delhi. pp. 107-123 (2009).
vstedal, D.O. and R.I.L. Smith: Addition and corrections to the lichens of Antarctica and
South Georgia. Cryptogamie, Mycologie. 25(4), 323-331 (2004).
vstedal, D.O. and R.I.L. Smith: Lichens of Antarctica and South Georgia. A guide to their
identification and ecology. Cambridge University Press, U.K. (2001).
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bryophytes: analysis of pressure-volume relationships. J. of Exp. Bot. 49, 1845-1854
(1998).
Stadelmann, E.J.: The derivation of the cell wall elasticity functions from the cell turgor
potential. J. of Exp. Bot. 35, 859-868 (1984).
Tuba, Z., M.C.F. Protor and Z. Csintalan: Ecophysiological responses of
homoiochlorophyllous and poikilochlorophyllous desiccation tolerant plants: a
comparison and an ecological perspective. Plant Growth Regulation. 24, 211-217 (1998).
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Chapter 9
* Eg225
mail: olegtro@aari.nw.ru, Fax:+7-812-352-2688, Phone: 7-812-337-3134
Arctic and Antarctic Research Institute, St.Petersburg, 199397, Russia
174
INTRODUCTION
Existing models of the atmospheric variability and change do not take into consideration
the short-term changes of solar activity. Indeed, the total energy, contributed by the solar
wind and the cosmic rays in the Earths atmosphere, is extremely insignificant in comparison
with the total solar irradiance. But, as distinct from the total solar irradiance, the energy of
solar wind and cosmic rays can increase in hundreds and more times in periods of high solar
activity. The attempts to find the cause-effect relations between the solar activity variations
and weather and climate changeability have a long story (Wilcox, 1975; Herman and
Goldberg, 1978). The galactic cosmic rays (GCR) altered by solar wind were usually
regarded as the most plausible agent of the solar activity influence on the Earths atmosphere.
The experimental data were presented showing the influence of the varying GCR flux on the
Earths weather and climate (Tinsley et al., 1989), on high cloud coverage (Pudovkin and
Veretenenko, 1995), on temperature in the polar troposphere (Pudovkin et al., 1996, 1997), on
the global total cloud cover (Svensmark and Friis-Christensen, 1997; Todd and Kniveton,
2001), on low cloud coverage (Marsh and Svensmark, 2003). These results suggest that just
cloudiness variation affected by cosmic rays lead to changes in the atmospheric and
meteorological characteristics.
However, the hypothesis about the determining influence of the galactic cosmic rays on
the cloudiness was not always supported by the subsequent, more detail research. It was
indicated that the correlation with GCR disappears when the cloud coverage is decomposed in
fractions by cloud type or height, by region (reduce for ocean basis), or by latitude (patterns
in the tropical zone are better associated with concurrent El-Nino) (Farrar, 2000). A
comprehensive study of the low cloud coverage for the last 120 years (Palle and Butler, 2002)
revealed that the global cloudiness increased during the past century regardless of variations
of GCR. The solar irradiance turned out to be correlated better and more consistently with
low cloud cover than the cosmic ray flux (Kristjansson et al., 2002). As a result, the
conclusion was maid that the mechanism linking the cosmic ray ionization and cloud
properties cannot be excluded, but its high efficiency is not obvious (Harrison and Carslaw,
2003).
At the same time it is well known that severe reductions in the galactic cosmic rays flux,
known as Forbush decrease (FD), are related to the disturbed, high speed solar wind,
emerging by the most intense solar flares. The disturbed solar wind is characterized by the
largest changes in such parameters as the solar wind pressure PSW = m n (VSW) 2 and the
interplanetary electric field ESW = VSW x BZ (or simply the southward component of the
interplanetary magnetic field (IMF BZS). The velocity VSW might vary with a factor of 2-3 at
maximum, whereas the IMF BZ components might change the sign and increase by some
factors of ten. The geoeffective solar wind parameters strongly affect (impact) the Earths
magnetosphere. It was noted that the FD beginnings at the Earths orbit are recorded
simultaneously with dramatic disturbances in the solar wind, and therefore, the atmospheric
effects, assigned to Forbush decreases, can be, in reality, influenced by the geoeffective solar
wind parameters (Troshichev et al., 2003).
Figure 1. shows, as an example, response of the cloudiness above Vostok station
(Antarctica) to Forbush decrease (left column) and to the IMF BZ minimum (right column)
during the winter season of 1974-1992 (Troshichev et al., 2008). The list of 24 Forbush
175
decreases was taken from the widely known analysis (Todd and Kniveton, 2001), but in our
case the Forbush decrease maximum has been used as a key date in the epoch superposition
method unlike to Todd and Kniveton (2001), who used the Forbush decrease beginning.
Indeed, in many cases it is difficult to determine the FD beginning unambiguously, as a result
the FD beginning dates sometimes are identified with as large a scatter as 5 days in various
studies. On contrast, the Forbush event maximum is easily and uniquely identified by
minimum in the galactic cosmic ray flux in each case.
The same list has been used to separate the IMF BZ minimum dates, related to the FD
events. Unfortunately, the IMF BZ data turned out to be available only for 15 events of 24. So,
the left column in Figure 1. is for the data of cloudiness above Vostok, allocated relative to
the FD maximum, whereas the right column is for the data allocated to the appropriate IMF
BZ minimum.
Figure 1. Behavior of the average Forbush decrease (FD), the average interplanetary magnetic field
(IMF) BZS component, and the appropriate cloudiness above Vostok for the most powerful FD events
during the winter season of 1974-1992 (list of Todd and Kniveton, [2001]). A key date (t = 0) was
taken as a day with FD maximum in the left column and a day with the IMF BZ minimum in the right
column (from Troshichev et al.,2008).
176
The results presented in the left column demonstrate that Forbush decrease coincides with
increased cloudiness, which starts three days ahead of the key date (FD minimum) and
reaches the 55 % maximum by the key date, the statistic significance being equal to 0.96. The
results presented in the right column demonstrate, with no less evidence, that cloudiness
above Vostok starts to increase one day before the IMF BZ minimum and reaches its
maximum next day after the key date. The statistical significance in this case is less, ss=0.91,
but we have to take into account that number of the available events was reduced in 1.5 times
while examining the Bz indicator instead of Forbush decrease. This example clearly shows
that changes of cloudiness may be successfully explained by Forbush decrease as well as by
the IMF variations.
177
and the corresponding warming in the ground layer, whereas the rise of the positive BZ
deviation is followed by the opposite reaction: the cloudiness decays and the cooling starts on
the ground layer. It should be reminded that Vostok is located at the ice dome at an altitude of
3.45 km above sea level, and, therefore, h=3.5 km corresponds to ground level at Vostok,
whereas h= 6 km corresponds to an altitude of ~ 2.5 km above the ice sheet level.
The results of the analysis (Troshichev et al., 2008), completed for conditions of the
negligible Forbush decrease, demonstrate that the sign of the BZ component defines the trend
of the cloudiness change (growth or decay), the cloudiness power being determined by the
value of the southward IMF component. It seems reasonable to suggest that the formation of
the cloud layer occurs at altitudes higher than ~ 5 km above the ice sheet.
Figure 2. Mean changes in cloudiness estimated from the radiation balance measurements (second
panel) and by the visual man-made observations (third panel), obtained for three gradations of the
negative deviation in the daily averaged IMF BZ component (-2<BZ< 1 nT, -2.5<BZ< 2 nT, BZ< 2.5
nT) during the winter season of 1974-1977 and 1985-1987 (from Troshichev et al., 2008).
178
Figure 3. Run of the daily averaged IMF BZ component for cases of strong negative and positive B Z
deviations (upper panel) and the appropriate effects in the cloudiness (second panel) and in temperature
at heights 3.5 km and 6 km above Vostok (third and forth panels) in years of the solar activity minimum
1974-1977 and 1985-1987 (from Troshichev et al., 2008).
179
Figure 4. Drainage pattern of near-surface katabatic winds (from Parish and Bromvich, 1991). Location
of the inner-continental stations Vostok, Dome C and South Pole is shown.
Figure 5. A conceptual scheme of the vertical mass circulation forced by the katabatic wind regime in
Antarctica (from Parish and Bromwich, 1991).
The spatial structure of katabatic winds is one of the most stable atmospheric phenomena
on the Earth (Schwerdtfeger, 1984). Phases of weak and strong katabatic winds alternate with
each other at regular intervals from few days (Egger, 1985; James, 1989) to 30-50 days
(Yasunari and Kodama, 1993). The model-based studies (Egger, 1985; James, 1989; Parish
and Bromvich, 1991; Parish, 1992) showed that the Antarctic katabatic wind regime appears
to be an important forcing mechanism for the circumpolar vortex around the periphery of the
Antarctic continent. As the experimental study of Yasunari and Kodama (1993) demonstrates,
the weak katabatic wind phase corresponds to a deep upper tropospheric circumpolar vortex,
whereas the strong katabatic wind phase corresponds to a weak circumpolar vortex. Thus, the
180
low atmosphere is not static above the Central Antarctic ridge: there is a powerful vertical
channel, where the air masses go down from upper troposphere to the Antarctic ice sheet.
181
between the key moment and the temperature change. At stations Vostok and Dome C a 15hours exposure affects the effective warming (up to T > +10) after 12 hours at a level of
statistical significance equal to 0.99.
However, the warming at station South Pole may be observed only under conditions of
very strong interplanetary electric field, and this link is not statistically significant. The
different effects in the ground temperature at stations Vostok and Dome C, on the one hand,
and South Pole, on the other hand, are easily explained by their different locations relative to
the katabatic wind system. Indeed, stations Vostok and Dome C are located on the Antarctic
ridge, which is the region of the descending tropospheric air mass flow, whereas South Pole is
located in the regions of the developed drainage stream.
As the results (Troshichev and Janzhura, 2004) showed, there is a weak tendency to
ground temperature decrease after the pressure pulses: the coolings at Vostok follow the large
pressure pulses within 2 hours and last about the next 24 hours (Figure 8). However, we have
to keep in mind that pressure pulses in the interplanetary shocks usually pass the Earth some
hours (from 0 to 12) ahead of the interplanetary electric field disturbances. So, the isolated
effects of the pressure increases can be seen, in principle, only whithin the first hours after the
pressure pulses. When the electric field comes into play, the influences of the pressure pulses
and interplanetary electric field would superpose on one another, and their combined effect in
the temperature deviations is observed.
Figure 6. Correlation between daily changes in ground temperature (T) at Vostok station (1978-1992)
and daily deviations in the interplanetary magnetic field BZ (upper panel) and in the geoeffective
electric ESW field (lower panel) (from Troshichev et al., 2003).
182
Figure 7. Character of temperature hourly changes T at stations Vostok, Dome C, and South Pole as a
function of number of the hourly interval with B Z <-2nT. Dotted line marks the key moment. Rate of
the warming is presented by dark signatures (from Troshichev and Janzhura, 2004).
183
Figure 8. Response of the ground temperature T at Vostok to pulses in the solar wind dynamic
pressure in 2000 2001, the key moment being determined by sharp increase in the dynamic pressure.
(from Troshichev and Janzhura, 2004).
It is meaningful that the response in temperature to the ESW influence is quite opposite in
the lower and upper troposphere. Figure 9. shows the profiles of the averaged daily
temperature deviations above Vostok station for negative and positive variations of ESW
(Troshichev et al., 2004). The temperature profile for the 1st day, preceding the zero day
(i.e. day of the maximum ESW deviation), is taken as the level of reference for all succeeding
days. The average warming at the ground level (h = 3.45-3.5 km) responds, within 1-2 days,
to the negative step in ESW, but at altitudes more than 10 km a cooling is observed (upper
panels). The opposite behaviour is typical of the positive step in ESW: the atmosphere gets
cooler at ground level, and gets warmer at h > 10 km (lower panels). Such regularity would
be observed if a cloud layer appears at altitudes 5-10 km under the influence of the negative
ESW variations, and disappears under the influence of the positive ESW variations.
184
Figure 9. Mean height profiles of the daily temperature deviations above Vostok under conditions of the
negative and positive ESW changes (from Troshichev et al., 2003).
185
at the first day and lowering to the ground level by the third day. In the context of the going
down air masses flow it implies that the interplanetary electric field effectively accelerates or
delays the descending tropospheric air masses.
Figure 10. Mean height profiles of the daily atmospheric pressure deviations above Vostok under
conditions of the negative and positive ESW changes (from Troshichev et al., 2004).
186
h= 15-25 km, 240-270 at h=10-15 km, and 210-240 at h=5-10 km, i.e. the horizontal winds
blow approximately toward the East.
To understand this regularity we have to remember that the masses falling vertically in
the rotating Earths system are deviated toward the East under the action of Corioles force.
Speed of this deviating motion is determined by formula ve = x r , where is the angle
speed of the Earths rotation ( = 1/240 /s), and r is radius-vector in the point of observation
(r ~ 880 km at latitude of Vostok station, = 82). Taking into account these quantities and
making allowance for angle ~ 8 between vectors and r, a rough measure of the deviation
speed is obtained, ve 10 m/s. It is just mean value of the horizontal speed that was observed
in aerological observations at altitudes 10-15 rm above Vostok station. Thus, we can suggest
that direction and speed of the regular wind in the surface atmosphere at a top of the Antarctic
ice dome is determined by deviating action of Corioles force on the descending air masses.
Figure 11. Dependence of the height profiles of the horizontal wind mean direction and mean speed
above Vostok on the negative ESW influence. The wind profiles for the -1st day preceding the ESW
impact (dotted lines) are taken as a level of reference (from Troshichev et al., 2004).
187
In response to the negative ESW leap the wind horizontal speed decreases in a key
(zero) day at altitudes higher 12 km and sharply increases at altitudes from 5 to 12 km (up
to 35 % at h = 8-9 km), the vertical wind profile for -1st day being taken as a level of
reference. The opposite regularity is typical of the positive ESW leaps. By the forth day the
wind speed recovers to the quiet level. Taking into account that Corioles acceleration should
be directly dependent on speed of the descending air masses, we suggest that increase or
decrease of the wind horizontal speed is indicative of acceleration or slowing-down of the
descending air masses. In such a case we came to conclusion that formation of cloud layer at
h=5-10 km is favourable to acceleration of the descending masses, whereas clear sky favours
their slowing-down. The mechanism of this linkage is unclear.
The acceleration (slowing-down) process should lead to the strong enhancement (or
reduction) of the atmospheric pressure just for day of the negative (positive) ESW leap, as we
have seen in Figure 10. It should be noted that the enhanced atmospheric pressure keeps
during some days in the surface level (~ 3.5 km at Vostok station). One might expect that
occurrence of the enhanced atmospheric pressure in the Central Antarctica will led to
violation of the wind regime above the whole Antarctica.
188
One can see from Figure 13. that occurrence of the anomalous winds at the coast stations
was preceded by the increase of southward IMF BZ and was delayed one-two days relative to
the wind strengthening at Vostok. Not all wind violations starting at the Antarctic Ridge are
extended to cover the overall coast area: only 82% of anomalous winds detected at Vostok
reach Casey, located in the Indian ocean sector and as few as 41% reach Neumayer, located in
the Atlantic sector.
Figure 12. Distribution of the regular (solid arrows) and anomalous (dashed arrows) winds at the
Antarctic stations (from Troshichev et al., 2008).
Figure 13. Anomalous winds at stations Vostok, Neumayer, Casey and Russkaya in their relation to
changes in the IMF BZ for winter seasons of 1981-1989, the day with anomalous wind occurrence at
each particular station being taken as a key date (from Troshichev et al., 2008).
189
It seems likely that extent of the coast station involvement in the process of the wind
violation varies from one event to other. The anomalous winds at the coast stations,
associated with high speed (V > 6m/s) anomalous wind at Vostok, were accompanied by the
sharp reduction of the westward wind speed and by the succeeding wind turning toward the
equator. This regularity corresponds to decay of the circumpolar vortex while increasing the
katabatic winds, noticed in (Yasunari and Kodama, 1993). The efficiency of the anomalous
winds influence on the circumpolar vortex turned out to be dependent on duration of the
southward IMF action: the 3-days influence of southward IMF with average BZ ~ 3 nT (10
events) preceded the anomalous winds, which were observed simultaneously at the all coast
stations.
190
been only included in the examination. The wind rose for the 4th month, preceding the ElNio onset, has been taken as a level of reference for all succeeding months, and the
appropriate differential wind roses, characterizing the changes in the wind azimuths for -3d,
-2d, -1st, and zero (El-Nio onset) months have been constructed. Results of the analysis
testified that the evident excess of winds above the level of reference was observed at angles
195-210 for Vostok, 185-215 for Russkaya, and 270-300 for Leningradskaya, which just
correspond to azimuths of the anomalous winds at these stations. As Figure 15. shows, the
anomalous winds are observed during -2d and -1st months, preceding the El-Nio onset, and
did not typical of -3th and zero months (as well as of succeeding months, which are not shown
in Figure 15).
Figure 14. Behavior of SOI index in course of large negative long-lived and short-lived SOI deviations
during 1868-2002 (from Troshichev et al., 2005).
191
Figure 15. Differential wind roses, characterizing the changes in wind occurrence in course of El-Nio
events for stations Vostok (a), and Russkaya (b) (from Troshichev et al., 2005).
Availability of statistically significant relationships between the disturbed solar wind and
anomalous winds above the Antarctica, on the one hand, and between the anomalous winds
and SOI, on the other hand, makes it possible to suggest that linkage between the disturbed
solar wind and development of SOI would be presented. Unfortunately, the monthly IMF data
are very incomplete for years preceding 1998, and the relation of SOI to AE index (instead
of IMF) was examined in (Troshichev et al., 2005). Only sharp SOI events, in which
the monthly value of SOI (negative or positive) was changed by 1 or more, have been
included in the analyses. The month of this sharp change has been taken as a zero date in the
superposition analysis.
Figure 16. shows behavior of SOI index for three types of changes in the southern
oscillation: (a) sharp declination of SOI with subsequent keeping of negative value during
many months (basically, it is real El-Nio events with sudden onset) observed in 1969, 1972,
1982, 1986, 1991, 1994, (b) sharp short-lived declination of SOI (1959, 1961, 1979, 1980,
1984, 1985), and (c) short-lived increase of SOI (1962, 1964, 1970, 1975). The low panel in
Figure 16 presents the proper changes in AE index. One can see, that in case of long negative
SOI deviations the mean magnetic activity starts to increase 2-3 months before the beginning
of El-Nio and reaches maximum just in month of the beginning (s.s. = 0.96).
192
Figure 16. Relationship between changes in monthly values of SOI and AE indices for long-lived
negative deviation of SOI (a), and short-lived negative and positive deviation of SOI (b) (from
Troshichev et al., 2005).
Figure 17. A conceptual sketch of generation of the electric currents in the low-latitude boundary layer,
while the solar wind coupling with the magnetosphere (adopted from Troshichev, 1982). It is shown the
magnetosphere dawn-dusk cross viewed from the Sun. The arrows denote the electric currents. The
field-aligned currents, flowing in the southern and northern polar caps at the dawn side and flowing out
at the dusk side, provide the polar cap electric voltage.
193
194
Figure 18. Diagram illustrating the global atmospheric electric circuit and the causes of its temporal and
spatial variation (from [Tinsley and Zhou, 2006]). The vertical scale is greatly exaggerated below 120
km and greatly compressed above.
CONCLUSION
Thus, the concept of the solar wind influence on processes in the Antarctic atmosphere
seems to be convincingly verified with use of all available meteorological and aerological
observations (Troshichev et al., 2003, 2004, 2005, 2008; Troshichev and Janzhura, 2004;
Troshichev, 2008). The disturbed solar wind has a greater impact on atmosphere processes in
the central Antarctica, where the large-scale system of vertical circulation is formed during
the winter seasons. The impact is realized through the interplanetary electric field ESW, which
influences the magnetosphere field-aligned currents, creating the appropriate polar cap
electric voltage. The ionospheric electric potential strongly affects the global electric circuit,
which is generated by the tropic thunderstorms and is closed mainly in polar caps.
Two processes are brought into operation when the IMF BZS component and the
appropriate electric field ESW strongly increase. The first of them is formation of the cloud
layer above the Antarctic Ridge, where the air masses income into the central Antarctica from
above troposphere. Under the usual conditions the atmosphere on the Antarctic ridge is in
state of the thermal quasi-equilibrium owing to superposition of the adiabatic warming of the
descending air masses and stable radiation cooling of air situated at the ice sheet. The cloud
layer will efficiently backscatter the long wavelength radiation going from the ice sheet, but it
will not affect the adiabatic warming process. As a result of the radiative cooling reduction,
the atmosphere should be warm below the cloud layer and would be cool above the layer. Just
such regularity is observed during specific events of the sudden warmings, happening from
time to time in the central Antarctica.
The second process is acceleration of the air masses incoming into the central Antarctica
from troposphere. This process causes the sharp increase of the atmospheric pressure in the
surface layer and gives rise to collapse of the regular large-scale wind system above the entire
195
Antarctica. The draining winds are strongly strengthened and the circumpolar vortex about
the periphery of the Antarctic continent correspondingly decays.
Both processes are strongly linked and lead to the cold air masses flow out to the
Southern ocean. The latter process evidently destroys the regular relationships between the
sea level pressure fluctuations in the Southeast Pacific high and the North AustralianIndonesian low and promote the El-Nio beginning.
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Tinsley, B.A. and C.W. Deen: Apparent tropospheric response to MeV-GeV particle flux
variations: a connection via eletrofreezing of supercooled water in high-level clouds? J.
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Tinsley, B.A. and R.A. Heelis: Correlations of atmospheric dynamics with solar activity:
evidence for a connection via the solar wind, atmospheric electricity, and cloud
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Tinsley, B.A. and L. Zhou: Initial results of a global circuit model with variable stratospheric
and tropospheric aerosols, J. Geophys. Res., 111, 16205-16223 doi:
10.1029/2005JD006988 (2006).
Todd, M. and D. Kniveton: Changes in cloud cover associated with Forbush decreases of
galactic cosmic rays, J. Geophys. Res., 106, 32031-32041 (2001).
Trenberth, K.E.: Planetary waves at 500 mb in the southern hemisphere. Mon. Weather Rev.,
108, 1378-1389 (1980).
Troshichev, O.A.: Polar magnetic disturbances and field-aligned currents. Space Sci.
Reviews., 32, 275-360 (1982).
Troshichev, O.A., L.V. Egorova and V.Ya. Vovk: Evidence for influence of the solar wind
variations on atmospheric temperature in the southern polar region. J. Atmos. Solar-Terr.
Phys., 65, 947-956 (2003).
197
Troshichev O.A., L.V. Egorova and V.Ya. Vovk: Influence of the solar wind variations on
atmospheric parameters in the southern polar region. Adv. Space Res., 34, 1824-1829
(2004).
Troshichev, O. and A. Janzhura: Temperature alterations on the Antarctic Ice sheet initiated
by the disturbed solar wind. J. Atmos. Solar-Terr. Phys., 66, 1159-1172 (2004).
Troshichev, O.A., L.V. Egorova and V.Ya. Vovk: Influence of the disturbed solar wind on
atmospheric processes in Antarctica and El-Nino Southern Oscillation. Mem. Soc.
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Troshichev, O., V. Vovk and L. Egorova: IMF associated cloudiness above near-pole station
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Chapter 10
1. INTRODUCTION
Antarctica has a fundamental role in the global climate system. However due to its
remoteness and inaccessibility, relatively little is known about the processes occurring in the
interior of the ice sheet. In the long Antarctic winter, with no short-wave radiation, the surface
is cooled continuously through negative net long-wave radiation which is enabled by clear
sky conditions and the very cold and dry overlying atmosphere. This energy loss is partially
compensated by the turbulent and sub-surface heat fluxes which extract heat from both the
atmosphere and the ice sheet, cooling the near surface air and snow. As a consequence of this
*
E-mail : s.argentini@isac.cnr.it
ISAC-CNR via del Fosso del Cavaliere, 100, 00133 Roma, Italy
200
radiative cooling strong surface-based inversions develop. Phillpot and Zillman (1970)
studied the climatology of the temperature inversion across the Antarctic continent, they
found that the inversions over South Pole are on average about 20 K in winter, while those
over the highest parts of the plateau are on average 25 K. Connolley (1996) gives an overview
of the temperature inversion at 21 Antarctic stations (whose 3 in the interior); following
Jouzel and Merlivat (1984) he derived a regression between the surface temperature and the
inversion strength all over the Antarctic continent also showing the dependence of this
relationship on the terrain slope. Other analyses were done to investigate the link between the
near-surface winds and the temperature inversions (Dalrymple et al., 1966 and Lettau and
Schwerdtfeger, 1967). Hudson and Brandt (2005) studied the surface-based inversion over the
Antarctic plateau; they focused on the measurements from South Pole and Dome C, but the
analyses were done for a short period and limited to the first 30 m of atmosphere at South
Pole, and restricted to the summer season at Dome C.
In presence of a surface slope, the denser and colder air masses of air adjacent to the
surface are forced down-slope due to the horizontal pressure gradient, and deflected to the left
by the Coriolis force. As a result the well known katabatic winds take place. The katabatic
wind regime can be occasionally interrupted by the horizontal advection of warm air masses,
or the presence of strong large-scale winds.
On the other hand, during the Antarctic summer, the absorption of short-wave radiation
introduces a diurnal cycle in the behaviour of the solar radiation. Although the amount of
solar radiation available to heat the surface is limited by the high surface albedo the surface
warming may cause a weak convection and the formation of a Convective Boundary Layer
(hereafter CBL) during daytime (Mastrantonio et al., 1999, Argentini et al., 2005).
Reliable measurements of the radiation balance at the surface on the Antarctic ice sheet
are important to assess its role as heat sink in the climate system of the earth, as ground truth
for satellite observations (e.g., to distinguish between clouds and the snow surface) and for
validation of atmospheric models (King and Connolley 1997). They may also be used to
develop albedo parameterizations of dry snow for atmospheric models or, for radiation
parameterization in energy and mass balance and ice dynamical models (Van de Wal and
Oerlemans 1997). A reliable estimate of the Surface Energy Balance (hereafter SEB) is not
possible without an accurate measurement of the radiation components (Bintanja and van den
Broeke 1995; Van As et al. 2005b) which can be affected by several problems related to snow
properties and sun elevation angle. Long-wave fluxes dominate the surface heat budget in
Antarctica because of the regions high solar albedo and prolonged periods of darkness.
The Antarctic atmosphere is highly transmissive for solar radiation, especially in the high
interior plateau where the atmosphere is thin and the concentrations of clouds, water vapour
and aerosol are low. Even in full summer, when the solar zenith angles are large, the finegrained, dry and clean snow surface absorbs only 525% of the incoming short-wave
radiation (Carroll and Fitch, 1981). On the other hand the snow, like most natural surfaces,
has a high long-wave emissivity (0.98) (Wiscombe and Warren, 1980) so that it effectively
looses heat in the form of long-wave radiation. In combination with an atmosphere that is
cold, dry, thin, clear and clean, this leads to a pronounced (long-wave and all-wave) radiation
deficit at the surface in winter. An average turbulent transport of sensible heat from
the atmosphere to the surface compensate for this heat loss. This loss of energy makes
the Antarctic ice sheet a major heat sink in the Earths atmosphere and introduces a strong
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 201
coupling between the radiation balance and near-surface climate (Dutton et al., 1991; Stanhill
and Cohen, 1997).
Despite the number of observational studies of SEB carried out in Antarctica, only few
measuring campaigns have been done during the winter and in the interior, due to the dearth
of occupied stations on the polar plateau and the enhanced difficulty of maintaining
instruments on robotic platforms. The above notwithstanding, the number of winter time
experimental campaigns in Antarctica during the winter is slowly increasing, although with
often unknown quality of data.
A short history of the SEB studies from unmanned stations is available, from Weller,
1980; Carroll, 1982; King and Turner, 1997; Wendler et al., 1988; Bintanja and van den
Broeke, 1995; Reijmer et al., 1999; Argentini et al., 2005; van As et al., 2005a. Some of these
studies evidenced that the magnitude of the surface heat fluxes exhibit strong spatial and
temporal variations, which reflect the differences in the prevailing meteorological conditions
and in the state of the surface. King et al. (2006) describe the summer SEB at two contrasting
Antarctic sites: the interior site of Dome C (75S 123E) at 3306 m a.s.l and the coastal site at
Halley (75S, 26W) at 30 m a.s.l. At both stations, the short wave radiation produces a
diurnal cycle in net radiation, which results in a diurnal cycle of surface temperature. Despite
this similarity, the summer-time SEB differs significantly at the two sites for what regards the
behaviour of the sensible heat flux. Convection signature is observed on acoustic radar
echograms at the cold Dome C site, while is absent at Halley. This is due to the large
summer-time net radiation at Halley generating melt, which limits the partitioning of the SEB
into convection.
Due to the extreme temperatures, dryness and the high altitude the Antarctic plateau is a
potential ideal site for astronomical observations otherwise possible only from space. For this
reason in the last years the optical turbulence over the Antarctic plateau has been object of
studies by astronomers (for ex. Hagelin, 2008), the optical turbulence intensity depending on
vertical gradients of the wind velocity and on the temperature inversion strength (Marks et al.,
1999).
A large atmospheric field experiment STABLEDC (Study of the STAble Boundary Layer
Environmental at Dome C) was held at the French-Italian station of Concordia located on the
Antarctic plateau at Dome C during 2004-2005. This was the first over wintering at
Concordia. The aim of the field experiment was to test an observing system to study the
processes occurring in the long-lived stable, and the weak convective atmospheric boundary
layers, observed during winter and summer respectively, and to collect the parameters
relevant for the atmospheric models. The results obtained during this experiment are shown in
the following sessions.
202
The climate consists of a synoptic coastal influence which brings relatively warm and cloudy
air masses, and gravity driven flow occurring under cold, clear, sky conditions. Low wind
speeds combined with frequent stable boundary layers and small surface roughness result
most of the time in low atmospheric thermal turbulence at Dome C.
The annual mean wind speed is 3.4 m s-1 with extreme values up to 16 m s-1. Monthly
mean temperatures were 39 C in summer and 61 C in winter. Most of the time, strong
surface inversions occur at Dome C, creating a large cold air source feeding the katabatic
winds observed in some zones of confluence along the East Antarctic coast.
In situ and ground based remote sensing sensors were used to monitor the behaviour of
the meteorological parameters. The instrumentation as well as the measurements done during
the field experiment are listed in Table 1.
Turbulence and radiation measurements were made in the period November 2004 January 2006. Turbulent heat and momentum fluxes were derived from eddy covariance
measurements (Lee et al., 2004) using a Metek USA-1 sonic thermo-anemometer sampling at
10 Hz installed 3.6 m above the snow surface; long- and short-wave radiation components
were measured using a Kipp and Zonen CNR-1 net radiation sensor system installed 1 m
above the snow surface. The heat flux within the snowpack was measured at a depth of 50
mm using a Campbell Scientific HFP01 heat flux plate. A mini-SODAR (Sound Detection
and Ranging) Doppler system (Mastrantonio et al., 1999) provided a continuous record of the
structure of the atmospheric boundary layer along the year. A passive Meteorological
Temperature Profiler (MTP-5P) (Kadygrov and Pick, 1998) was used for the remote
measurement of the air temperature profile.
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 203
The instrumentation was placed approximately 1000 m South of the main base buildings.
Since the dominant wind direction is from the sector 150-210 this ensured an unobstructed
fetch over the measurement site.
In addition to the measurements described above, standard meteorological parameters provided by the AWS (Automatic Weather Stations), and radiosonde profiles (once a day) were available from other projects.
Table 1. Sensor and measured micrometeorological variable
SENSOR
Radiometer mod.CNR-1 (Kipp and
Zonen) with
two pyranometers (CM3) up and down
two pyrgeometers (CG3 ) up and down
Conventional HFP01 heat flux plates
0, 5, 15, 30, 50 cm
A sonic anemo-thermometer mod.
USA-1 (Metek) and a fast response
LICOR Lyman-alpha-hygrometer (only
summer )
13-m Tower : thermometers,
hygrometers and wind probes at 1.25,
2.5, 5, 10 and 13 m
A triaxial Doppler mini-SODAR
Range 12 - 400 m , Resolution 13 m
Micro-lidar 532 nm wavelength , Range
300 m
Passive Microwave radiometer MTP5P by Kipp and Zonen. Range 0-600 m
Radio soundings
MEASURE
Radiative budget: - Incoming and outgoing
short-wave and long-wave radiation.
-Net Radiation
-Albedo
Sub-surface energy fluxes, snow temperature
profiles:
-Snow heat fluxes
Energy budget:
-Turbulent Fluxes (Heat, Latent, Momentum)
For convenience, all subsequent results are presented with reference to local time. The
terms night-time and nocturnal are referred to the periods of negative surface radiation
balance.
204
The annual mean wind speed is 4.4 ms-1 with extreme values up to 12 ms-1. The mean
annual temperature is 45 oC. The lowest temperatures (-72 C was the minimum) are
observed during the winter while the highest temperatures (-29 C was the maximum) are
observed during the summer.
Table 2. Mean values of the wind speed, temperature and sensible heat flux
for the measurement periods: summer, autumn, winter and spring
Period
All
Summer
Autumn
Winter
Spring
Summer
Parameter
2004/12/14
to
2006/01/13
2004/12/14
to
2005/01/31
2005/02/01
to
2005/03/31
2005/04/01
to
2005/09/14
2005/09/15
to
2005/11/14
2005/11/15
to
2006/01/13
Mean wind
speed (m/s)
4.4 2.0
3.5 1.6
4.2 1.7
4.9 2.0
4.8 2.2
4.0 2.0
Mean
temperature
(C)
-44.8
12.2
-29.3 4.7
-47.2 7.8
-54.8 6.9
-46.8 7.6
-30.2 5.7
Mean
sensible
heat flux
(Wm-2)
-4.5 10.4
3.2 8.9
-4.8 8.1
-10.6 8.8
-6.1 9.5
2.4 9.0
Strong long-lived ground based inversions occur most of the time at Dome C with the
exception of the summer days during the hours of maximum insulation (and positive values
of net radiation). These inversions contribute to create a large cold air source producing and
feeding the katabatic winds observed in most of the glaciers confluence zone along the East
Antarctic coast.
Few clouds are generally present in the sky above Dome C since cloud cover and
precipitation decrease as one moves inland from the coast, and the frequency of occurrence of
active weather systems is low (King and Turner, 1997). However Argentini et al. (2001) have
shown that warming events are periodically observed at Dome C during the winter; during
these periods the surface temperatures sometimes may reach the summer values. In
correspondence of the warming events the wind direction changes from 180 (the most
frequent wind direction) to 0 (which is from the coast), indicating that the warming events
are correlated to phenomena originating along the coast (i.e. advection of warm air).
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 205
30 %
330
30
24 %
18 %
300
60
12 %
6%
0 v < 2 ms -1
90
270
2 v < 4 ms -1
4 v < 6 ms -1
240
120
v 6 ms -1
210
150
180
The wind speed during the summer (Figure 4.a) exhibits a clear diurnal variation with a
peak during the warmest hours of the day. As shown also by King et al. 2006, a little variation
in wind speed occurs between 1900 and 0600 local time (LT). After 0600 LT, the wind speed
increases gradually until local noon, when it is about 1.5 ms-1 higher than it was at 0600 LT.
The wind speed then remains fairly constant until 1500 LT, after which it falls off at a rate
somewhat faster than it increased during the morning, reaching its night-time value at
around 1900 LT.
206
Speed (ms-1)
9
8
7
6
5
4
3
2
1
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan
Figure 3. Wind velocity behaviour at Concordia station from January 2005 to January 2006.
Figure 4. a-d. shows the diurnal behaviour of the wind speed for summer (a), autumn (b),
winter (c) and spring (d). The diurnal variations observed at Dome C during the summer are
typical of those seen at locations where a convective boundary layer develops in response to
daytime heating. During the autumn the wind velocity is fairly constant around 4.2 ms-1
(Figure 4.b.), 5 ms-1 in winter (Figure 4.c.), and 4.8 ms-1 in spring (Figure 4.d.).
4.2. Temperature
The temperature histogram in the range -70 C / -20 C at steps of 5 C is shown in
Figure 5. The ordinate axis gives the percentage of temperature values observed in a given
temperature range. This frequency distribution shows two peaks. One peak is centred at -50
C, the other between -35 C and -20 C. The peak at -50 C corresponds to winter, spring
and autumn temperatures. The secondary peak includes instead the summer values.
Figure 6. shows the daily average of the 3-meters AWS (Automatic Weather Station)
temperature. The temperature has a strong seasonal cycle with values varying between -20
C, during the short summer, and -70 C in the long and coreless winter. Period of warming
events are evidenced during the winter, when the temperature in few cases may reach values
typical of summer.
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 207
6
15/11 - 01/02 ( SUMMER )
(a)
5
4
3
6
(b)
5
4
3
6
01/04 - 15/09 ( WINTER )
(c)
5
4
3
6
15/09 - 15/11 ( SPRING )
(d)
5
4
3
10
11
12
13
14
15
16
17
18
19
20
21
Time (Hour)
Figure 4. Average diurnal cycles of the wind (a) summer, (b) autumn, (c) winter, (d) spring, at
Concordia station from January 2005 to January 2006.
16
14
Occurrence (%)
12
10
-70
-65
-60
-55
-50
-45
-40
-35
-30
-25
Temperature (C)
Figure 5. Temperature histogram at Concordia station from January 2005 to January 2006.
-20
22
23
208
-10
Temperature (C)
-20
-30
-40
-50
-60
-70
-80
N D
F M A M J J A S O N D
Time (Months)
Figure 6. Temperature behaviour at Concordia station from January 2005 to January 2006.
The warming events last for about 5 days and have a periodicity of about 15 days. During
these periods the temperature may increase of several degrees (Argentini et al., 2001).
Carroll, 1982, Stone et al. 1990, Stone and Kahl, 1991 and Stone, 1993 on a study at
South Pole evidenced that these warming events were mostly observed in presence of
cloudiness. Neff (1999) analysing the periods of cooling and warming at South Pole
correlated these phenomena to the variations in wind direction in the elevated layer toward
Nord-East, sometimes this variation was accompanied by cloud advection from the West or
from the Weddel sea. Carrol (1982) suggested two possible mechanisms: advection of warm
air and/or vertical mixing of air from different layers. Othake (1978) analysed the particles
trajectory across Antarctica; he showed that the warming was mostly due to the intrusion of
warm and moist air and to the condensation of nuclei which originate from the Wedding sea
and reach the pole producing a variety of clouds type. Schwerdtfeger and Weller (1977)
related the surface warming to the variation of long-wave radiation emitted by clouds
associated at moist air from the upper part of atmosphere.
Figure 7.a-d. show the mean diurnal variation of the temperature at Dome C for summer
(Figure 7.a.), autumn (Figure 7.b.), winter (Figure 7.c.), spring (Figure 7.d.). The sonic
temperature behaviour is similar in summer, autumn and spring, with the peak of the
temperatures 23 hours after the local noon. The amplitude of the diurnal temperature
variation is ~10C during the summer (the maximum and minimum temperatures are
respectively -25C and -35C) and ~5C during the spring and autumn (maximum
temperature -45C, minimum temperature -50C ). During the winter the average value of the
temperature is ~ -55C.
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 209
-20
15/11 - 01/02 ( SUMMER )
-25
(a)
-30
-35
-40
Temperature (C)
-40
01/02 - 01/04 ( AUTUMN )
-45
(b)
-50
-55
-60
-40
01/04 - 15/09 ( WINTER )
-45
(c)
-50
-55
-60
-40
15/09 - 15/11 ( SPRING )
-45
(d)
-50
-55
-60
10
11
12
13
14
15
16
17
18
19
20
21
22
23
Time (Hour)
Figure 7. Average diurnal cycles of the sonic temperature (a) summer, (b) autumn, (c) winter, (d)
spring, Concordia station from January 2005 to January 2006.
Figure 8. shows the profiles for summer (a), autumn (b), winter (c) and spring (d) in the
vertical range 0-300 m. Each figure contains three profiles corresponding at averages over
different hours of the day. A diurnal profile (averages between 1000 - 1400 LT), an all
day profile, and a nocturnal profile (averages between 2200-0200 LT). An unstable profile
is observed in summer when the sun elevation reaches a peak. For all the other hours/seasons
a stable boundary layer is observed. The temperature increases of about 5 C in the first 100
meters during the summer and 20 C during the winter.
The result of a detailed analysis of the potential temperature gradients over all the winter
is given in Figure 9. The graph shows the density contour plots of potential temperature
gradients in the first 150 m of atmosphere. It is evident that the temperature gradient generally
ranges between 0.1 and 1 (C/m). However the strongest gradients occur below 40 m. This
result is in general agreement with the observations by Agabi et al., 2006.
210
300
250
Diurnal
Mean
Nocturnal
a)
250
Height (m)
Height (m)
150
100
50
50
0
-40
-35
-30
-25
Temperature (C)
0
-60
-20
300
-55
-50
-45
-40
Temperature (C)
-35
-30
300
Diurnal
Mean
Nocturnal
c)
250
200
Diurnal
Mean
Nocturnal
d)
200
Height (m)
Height (m)
150
100
150
150
100
100
50
50
0
-70
b)
200
200
250
Diurnal
Mean
Nocturnal
-65
-60
-55
-50
Temperature (C)
-45
-40
0
-60
-55
-50
-45
-40
Temperature (C)
-35
-30
Figure 8. Temperature profiles diurnal (crossed line), averaged (black dotted line), nocturnal (black
dotted line), during a) Summer, b) Autumn, c) Winter, d) Spring at Concordia station at Dome C from
January 2005 to January 2006.
Figure 9. Iso-Colour zone graphic of potential temperature gradient during the 2005 winter at Dome C.
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 211
H0 (Wm-2)
5
0
-5
-10
-15
-20
-25
-30
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan
Figure 10. Sensible heat flux behaviour at Concordia station at Dome C from January 2005 to January
2006.
Rnet H 0 H L G 0
(1)
where Rnet is the net radiation, H0 is the sensible heat flux, HL is the latent heat flux and G is
the conductive heat flux through the snowpack. We use the sign convention that fluxes of
212
energy directed towards the snow surface are positive. Rnet may be broken down into its
components
Rnet SW SW LW LW = SWnet+LWnet
(2)
where SW is the incoming short wave radiation, SW is the reflected short-wave radiation,
LW is the incoming long-wave radiation, LW is the outgoing long-wave radiation, SWnet
and LWnet are the net short wave and the net long wave respectively. Careful quality control
of these radiation data is essential due to the large absolute values generated by the
instruments, from which a relatively small difference is extracted.
The annual behaviour of the daily mean of the radiative budget components and the
Radiation at Top Of Atmosphere (RTOA) for 2005 are shown in Figure 11. As expected an
annual variation and a diurnal variability are present. The SW reaches the peak value during
the summer (480 Wm2 ) and decreases to 50 Wm2 at the beginning of the winter.
600
RTOA
-2
Radiation (W m )
500
400
SW
SW
LW
LW
300
200
100
0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Local Time (day)
Figure 11. Time series of daily mean of RTOA (black line), SW (--), SW (dot), LW (--), LW
(circle) at Concordia station from January 2005 to January 2006.
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 213
100
SWnet
LWnet
Rnet
80
-2
Radiation (W m )
60
40
20
0
-20
-40
-60
-80
-100
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Local Time (month)
Figure 12. Time series of daily mean SWnet (dotted line), LWnet (star) and Rnet (circle) at Concordia
station from January 2005 to January 2006.
The LW ranges between 200 Wm2 (in summer) and 80 Wm2 in winter. In winter during
cloudy days the LW increases of 20-30 Wm2. The seasonal variation of LW reflects the
cycle of the surface temperature with maximum values (~ 200 Wm2) in summer and
minimum values (~ 100 Wm2) in winter. During the summer the short-wave radiation
absorption helps in reducing the intensity of the surface inversion, instead, during the winter,
surface based inversion may also decrease during the warming events as the surface
temperature increases. Figure 12. shows time series of SWnet, LWnet, and Rnet. The LWnet is
the only component contributing to the net radiation during the winter. In summer LWnet and
SWnet have opposite sign, as a result Rnet (20 Wm2), although small, is positive. For each
season the diurnal cycle of the four components of the radiative budget is shown. The diurnal
cycles of SW is shown in Figure 13. The SW peaks around local noon. During the summer
the peak is about 800 W m-2. In spring and autumn the peak value reduces to 400 W m-2 while
in the full winter night the SW is zero.
The diurnal variation of incoming long-wave radiation at Dome C is shown in Figure 14
a-d. A clear diurnal cycle is apparent at Dome C in the outgoing solar radiation for all seasons
with the exception of the winter. This suggests that there is a diurnal cycle in temperature in
the lower part of the atmospheric column at Dome C as well as at the surface. In summer such
diurnal variability is consistent with our hypothesis of a diurnal CBL at Dome C. Outgoing
long-wave radiation is simply related to snow surface temperature, Ts, through Stefans law
LW Ts 4 (1 ) LW
(3)
where is the long-wave emissivity of the snow surface and is Stefans constant. Since
is approximately constant and has a value close to unity, the first term on the right hand side
of (5) dominates and variations in LW closely follow those in Ts. The diurnal variation of
LW during the summer reflects the larger diurnal range in near-surface air temperature at
Dome C. In the other seasons such variation it is not observed. Figure 15. a-d illustrate the
diurnal variation of Rnet obtained by eq. 2 at Dome C. During the night time, Rnet at Dome
C is typically -50 W m-2.
214
1200
15/11 - 01/02 ( SUMMER )
(a)
800
400
0
600
01/02 - 01/04 ( AUTUMN )
(b)
400
200
0
600
01/04 - 15/09 ( WINTER )
(c)
400
200
0
600
15/09 - 15/11 ( SPRING )
400
(d)
200
0
0
10
11
12
13
14
15
16
17
18
19
20
21
22
23
Time (Hour)
Figure 13. Shortwave daily radiation down for (a) summer, (b) autumn, (c) winter and (d) spring at
Concordia station from January 2005 to January 2006.
300
LW
(a)
LW
200
100
0
300
01/02 - 01/04 ( AUTUMN )
(b)
200
100
0
300
01/04 - 15/09 ( WINTER )
(c)
200
100
0
300
15/09 - 15/11 ( SPRING )
(d)
200
100
0
10
11
12
13
14
15
16
17
18
19
20
21
22
23
Time (Hour)
Figure 14. Longwave radiation up and down for (a) summer, (b) autumn, (c) winter and (d) spring at
Concordia station from January 2005 to January 2006.
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 215
100
15/11 - 01/02 ( SUMMER )
50
(a)
0
-50
-100
100
01/02 - 01/04 ( AUTUMN )
50
(b)
0
-50
-100
100
01/04 - 15/09 ( WINTER )
50
(c)
0
-50
-100
100
15/09 - 15/11 ( SPRING )
50
(d)
0
-50
-100
10
11
12
13
14
15
16
17
18
19
20
21
22
23
Time (Hour)
Figure 15. Diurnal cycle of net radiation for a) summer, b) autumn, c) summer, d) spring at Concordia
station from January 2005 to January 2006.
A regular diurnal cycle is observed for all the seasons with the exception of the winter
when there is not solar radiation. The peak of Rnet is about 80 W m-2 and 50 W m-2 during
the spring and autumn respectively.
216
solar radiation reaching the surface, consequently its depth varies from November to end of
February depending on the solar elevation angle. Figure 16.b shows the 1-hour averaged
temperature profile for the same day of Figure 16.a at 1400 LT. A weak convective activity is
observed close to the ground with a strong temperature inversion above this confirming the
mini-SODAR observations. By contrast, mini-SODAR records during the winter show no
echoes during the all day. The only exception is given by the days in which the arrival of
perturbations produce a thermal fluctuation that is strong enough to be revealed by the miniSODAR. Figure 17.a gives as an example of the thermal structure for 21 July, 2005, no
echoes are present during the entire day. Again if some echoes exists they are confined below
the first mini-SODAR range gate. Some field experiments carried out by astronomers
interested in site testing to the estimate the atmospheric optical seeing at Dome (Agabi et
al., 2006, Aristidi et al., 2005) have shown a poor ground seeing mainly due to a strong
turbulent boundary layer confined in the first 20 m 30 m close to the ground. In the future
we then plan to set the mini-SODAR in a configuration which will allow to study the
structure and depth of this shallow layer close to the ground.
600
b)
500
Height (m)
400
300
200
100
0
-25
-24.5
-24
-23.5
-23
Temperature (C)
-22.5
-22
Figure 16. (a) Thermal structure of the atmosphere during summer as seen by mini-SODAR for day 7
January 2005, (b) temperature profile by MTP-5P for the same day at 1400 LT.
Atmospheric Observations at Dome C, Antarctic Plateau, One of the Coldest Place 217
600
b)
500
Height (m)
400
300
200
100
0
-70
-65
-60
-55
-50
Temperature (C)
-45
-40
Figure 17. (a) Thermal structure of the atmosphere during winter as seen by a mini-SODAR for day 21
July 2005, (b) temperature profile by MTP-5P for the same day at 1400 LT.
Figure 17.b. shows the 1-hour averaged temperature profile for the same day of Figure
8.a at 1400 LT. A strong ground-based inversion of the order of 0.125 C/m is observed.
ACKNOWLEDGMENTS
This research was supported by the Piano Nazionale Ricerche in Antartide (PNRA) in the
frame of French-Italian Dome C project. The authors would like to thank the logistics staff
at Concordia for their support during the experimental fieldwork and all people which
contributed to the field experiments. A special thank to Ing. Guillaume Dargaud
overwintering at Concordia station during 2005, and to Sig. A. Conidi which participated to
the summer field operations during STABLEDC.
The authors also wish to thanks Dr. G. Mastrantonio, Dr. A. Viola, and Dr. Igor Petenko
which contributed to some of the results showed in these papers, for useful comments, and all
the work which has been done in the realization of STABLEDC.
218
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Publications of the Astronomical Society of the Pacific., 118(840), 344-348 (2006).
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characteristics of East Antartica meteorological parameters during 1994. J. Geophys.
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Argentini, S., A. Viola, A. Sempreviva and I. Petenko: Summer boundary-layer height at the
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Aristidi, E., A. Agabi, E. Fossat, M. Azouit, F. Martin, T. Sadibekova, T. Travouillon, J.
Vernin and A. Ziad: Site testing in summer at Dome C, Antarctica. Astronomy and
Astrophysics., 444(2), 651-659 (2005).
Bintanja, R. and M.R. van den Broeke: The Surface-Energy Balance of Antarctic Snow And
Blue Ice. J. Appl. Meteorol., 34, 902-926 (1995).
Carroll, J. J. and B.W. Fitch: Effects Of Solar Elevation And Cloudiness On Snow Albedo At
The South-Pole. J. Geophysical Res., 86, 5271-5276 (1981).
Carroll, J. J.: Long-term means and short-term variability of the surface energy balance
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Connolley, W.M.: The Antarctic temperature inversion. Int. J. Climatol., 16, 1333-1342
(1996).
Dalrymple, P. C.: A physical climatology of the Antarctic Plateau. Studies in Antarctic
Meteorology, (Eds. M.J. Rubin), Antarctic Research Series, 9, Amer. Geophys. Union.,
195-231 (1966).
Dutton E.G., R.S. Stone D.W. Nelson and B.G. Mendoca: Recent interannual variations in
solar radiation, cloudiness and surface temperature at the South Pole. J. Clim., 4, 848-858
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Hagelin, S., E. Masciadri, F. Lescaux and J. Stoesz: Comparison of the atmosphere above
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Hudson, S.R. and R.E. Brandt: A look at the surface based temperature inversion on the
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Jouzel, J. and L. Merlivat: Deuterium and Oxygen 18 in precipitation modelling of the
isotopic effects during snow formation. J. Geophys. Res., 89, 11749-11757 (1984).
Kadygrov, E.N. and D.R. Pick: The potential for temperature retrieval from an angularscanning single-channel microwave radiometer and some comparisons with in situ
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King, J.C. and W.M. Connolley: Validation of the surface energy balance over the Antarctic
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Chapter 11
222
INTRODUCTION
Global warming and ice melt is directly proportional to the rate of carbon burn of agrocellulose residues (ACR), coal and petroleum, partially differentiated in case of developing
countries and developed nations. Freon and fossil fuel consumptions, perhaps to be
considered more seriously, 1000 X time intensified, compared to the emission of CO2 caused
by burning of coal and ACR. Due to increasing carbon dioxide (CO2) emission, the physicochemical properties of stratosphere, amplifying the stratospheric reflections and solar (sun)
rays heat. This is equally being important in case of GHGs (CH4, CO, N2 and CO2) effects
in the history of mankind. The same phenomenon has happened on ancient earth, about 3.7
billions of years ago during Golden age of Reptiles, and caused the extinction of reptiles.
Ice changes to liquid water, if warmed above 4oC to its melting point and Vis- a Vis
changed to ice, if cooled to its freezing point. Earth's Polar Regions consists largely of ice and
exhibit dramatic change in temperature.
An increase of 0.1oC in temperature resulting from global warming may affect drastically
the extent of polar ice and the habitats of polar region. However this will not affect the change
of sea level, since the floating of ice follows the Archimedes principle, corresponding to the
displacement of water. The melt of continental ice (such as glaciers and ice sheets) raises sea
level. This happens because, ice on land when melts adds its water into the ocean which was
not there before. This extra water raises sea level.
For example, if the ice of Himalaya starts to melt it will affect the change of sea level of
bay of Bengal and Indian ocean and subsequently the same will affect the increase of water
level of Ganges and Bramhaputra belt and subsequenly may cause flood and the increase of
wastewaters, compared to potable drinking water. Although the melting of ice doesn't
significantly affect the sea level, if floats, there are other consequences which have to be
considered. Ice melt may cause the variations in salinity and temperature drive, global ocean
circulations, because the density of ocean water differs. Fresh water is less dense than salt
water and warm water is less dense than cold water. This thermohaline circulation may
sometimes refer the great ocean "conveyor belt" because it is one of earth's main mechanisms
for transporting energy (Edgerton, 2008).
The formation of sea ice, which is primarily made of pure water, leaves behind salt in the
water beneath the ice, resulting saltier and colder water that sinks through the water below it,
and thereby promoting the circulation. When ice melts, it adds fresh water to the ocean,
decreasing salinity and affecting the circulation pattern.
Any change to ocean circulation could have damaged the water cycle and weather
patterns. In addition, a change in the temperature or salinity of ocean water disrupts the
habitats and could have harmful effects on marine life forms that are sensitive to such
changes. The loss of sea ice in Polar Regions also threatens the survival of certain species,
such as the polar bear, which depend on the ice for hunting and breeding. Increasing
population will cause the increasing global warming due to higher CO2 emissions. Socioengineering (SE) application shows its significant application to control population and global
warming, in third world developing countries.
Application of Microbial engineering (ME) and microbes may play a significant role to
change the social impact in pollution abatement. So the reform of social economical
structure in third world in support of ME (microbial Engineering) may change the
223
engineering concept of sociology, the exploitation of natural resources in an appropriate, ecofriendly and cost-effective applications.
Such application may reduce the global warming. One example would be the reform of
water resources, and their recycling, which is exponentially growing as basic demand, with
increasing population. Although ME and SE are existing in two different boundaries, have
shown their common interest in technological innovations to optimize the demand of food,
health and shelter, the entropy (S) and chaos (Brahma and Basu, 2008). As per the
"Darwins Laws of Evolution, mankind belongs to the highest group of "Animal Kingdom"
adapting to all kinds of environmental factors in the course of 50 billions of years in early
earth evolution.
The best example may be giving by The Golden Age of Reptiles" and their extinctions.
This is to be considered seriously with the present global warming, ozone depletion,
population and pollutions. Since food, shelter, health and biological essential demands are
predominating. In addition to competition for better life, with material gains and
consumptions the increasing population proportionally relates the increasing pollution.
In this case the SE - application the reduction of population perhaps would be the best
solutions to resolve the problem of present world crisis of global warming. SE and
technological innovations essentially affect independent to color, race, religion, and
population control. This will ultimately be imposed by Enforced appropriate technology
(EAT) for specific technical applications Zero Effluent Discharge (ZED), waste to energy
(WTE), Microbial Enhanced Technology (MET) perhaps would integrate SE along with ME.
The global population and pollution will ultimately cross the boundaries of Life and
engineering Interfacial Phenomena, where animate and inanimate reactions will predominate
and perhaps will show the guideline to explain how the hybrid technology and innovations
will be essential for clean environment. Clean water, air and soil will be essential to utilize
Time and Energy as Human Factor (Brahma, 2004; 2007).
224
225
experienced warming and cooling cycles, roughly every hundred thousand years due
to these orbital shifts, but such changes have occurred over the span of several
centuries. Today's changes have taken place over the past hundred years or less.
(k) Other recent research has suggested that the effects of variations in the sun's output
are "negligible" as a factor in warming, but other, more complicated solar
mechanisms could possibly play a role.
ALTERNATIVE CONCEPT
Global warming is caused by green house gases (GHGs), which traps the suns infrared
rays in the earths atmosphere, which in turn heat up the earths atmosphere. This green house
effect warming is called as Global warming. The effects of green house are visible more
prominently in the recent years, with number of natural calamities on the rise in the whole
226
world. The global warming has happened in the past few years and is evident from the rise in
mean temperature of the earths atmosphere. The main causes for the global warming are
attributed to release of green house gases by human activities. The main gases contributing to
green house effect are carbon dioxide, water vapor, methane and nitrous oxide. The largest
producers of these gases are the thermal power plants, which burn the fossil fuels and produce
these gases in large quantities. The second biggest sources of these green house gases are the
road vehicles and industries (Archer, 2006; Brahma and Basu, 2008). The global warming has
led to an increase in mean earth surface temperature and thus melting of polar ice. There are
frequent melt down of glaciers that result in floods and other natural calamities. The melting
of polar ice had lead to increase the mean sea level. And further increase in temperature may
further melt the ice and lead to further increase in mean sea level, which will engulf low lands
of the continents. The effect of global warming is very important for animal kingdom. Some
animals have become extinct due to loss of their natural habitat or their inability to evolve to
the rapid changes in the climate. Also there is a change in their life style because of the
changes in the seasons. The migrating birds have changed their time of travel and also their
place of migration. The effect of global warming can be seen on seasons too. There is shift in
season cycle, as the summers are getting longer than the winters. This has affected the
animals and made them to change their lifestyle accordingly, and those who failed to do so
are perished or on the verge of extinction. The global warming is also responsible for the
introduction of some new diseases. The bacteria are more effective and multiply much faster
in warmer temperatures compared to cold temperatures. The same concept has been
introduced by the author in the form microbial engineering (ME), affecting the environment
and health. The increase in temperature has led to increase in the microbes that cause
diseases; subsequently the survival attitudes of microbes from ambient temperature to
mesophilic and to finally to thermophilic are increasing causing thereby pollution. Global
warming is also affecting the crop production. The sudden change in temperatures or sudden
set of rains destroys the crops productions. Also the flash floods and other natural calamities
affect the crop. As a matter of fact, global warming changes the earths atmosphere and is
getting more unpredictable rainfalls in areas, where the scanty of rainfall or drought were
predominated. The annual rainfall pattern may also be affected by Global Warming.
227
present times. GHGs are the ingredients of the atmosphere that add to the
greenhouse effect.
(b) 0.7-meter increase of sea level may act as a problem of environmental adaptations,
affecting directly to the ocean plankton, flora and fauna. Adaptation has an adverse
to biodiversity. CO2, Freon and Chlorofluorocarbons are all contributing the
disbalance of ecosystems effects and global warming. Environmental Protection
Agency (EPA); analyze the reports of Global climate changes. Research Resources
Defense Council indicates, that in their action plan, the sea rise and climate change
may be caused due to rapid change of global warming and the change of
Stratospheric Ozone structures. The same study was included in Washington Water
Zone Management Program, where they have marked the potential threat areas of the
earth surface.
Figure 1 .
Figure 2. a.
Figure 2. b.
Figure 1. Figure 2. (a.b.) represents the sectors, which influence percentile the global warming, caused
by various types of fossil fuel burns and the increasing propagation of warming with increasing the
years. Figure 2.b. shows the separate effect of Greenhouse Gases.
228
kilometer. GHGs like CH4, NOx, CO and CO2 may increase the effect. Among them CO2
may be considered as major culprit and may cause stratospheric hole as shown in Figure 3.
Once the sufficient solar rays enters the earth atmosphere, automatically by the theory of
light it will try to reflect, and be amplified in the cover of CO2 emitted GHG and will reflect
back to the earth surface, causing Global Warming.
The same will continue to change the entropy. Greeneries and plantations are prone to
absorbed the sun ray, photosynthetically to make their food in the form of glucose and
ultimately to cellulose, liberating at the same O2. This O2 is converting to O3 (Ozone) on the
surface of ocean in presence of sunlight, that absorbs IR (Infrared) and thereby the reduction
of thermal wave.
However in case of GHG and CO2 this IR could perhaps be captured by CO2 to protect
the reactions of O3 and reduce the photosynthetic activities of planktons, flora and fauna.
Therefore it is essential, that greeneries and increase of plantation may capture CO2.
Moreover sea planktons, algae and microbes are involved to capture CO2 as part of their
biosynthetic process.
Since there are variations of population on the surface of earth, CO2 emission may also
vary accordingly i.e., CO2 emits form carbon (in fossil fuel) burning, might be different
compared to CO2 emits by the burns of coal and ACR residues as fine particulates (Fick,
2004).
This is a type of formulation to describe the increasing of entropy due to stratospheric
hole and amplifications of sun-light as source of thermal energy, characterized by S = Q/
T. So the capturing of solar energy could be variable on the surface of the earth and may be
influenced by CO2 emission. Alternative concept could be the increasing population and
survivals demand.
More population means more pollution, which automatically generate CO2 emission.
Compared to developed nations, developing nations are consuming less energy, i.e., less
consumptions and burns of fossil fuel.
However they commit more CO2 emission, since they burn coal and ACR as direct
source of their survivals and energy resources, generating at the same time, nano- and pico
particles of CO2. From the part of population dense, it is also true that third world countries
will suffer from various types of environmental hazards and global warming, compared to
that of developed nations (Hasselmann, 1997).
Developed nations are basically controlled by low dense population, free of pollution and
cold atmosphere. So statistically even if they consume more energy, by burning fossil fuel,
they are in position to emit less CO2 compared to developing thick populated nations. It might
be true that to control cold, the developed nation burns coal in thermal power stations and use
strategic measure to heat their automobiles, if compared to third world nations, where Freons
are used for cooling, emits CO2 , 10,00 times more than fossil fuel burn. This relies the change
of entropy (Levi et.al., 1992).
Entropy will influence the change of biological system, chaos in biodiversity and
automatically the danger for the extinction of living system. Markovn relation; spatial
propagation of Markov truncated electrostatic interactions, existing between one amino acid
and the others at the protein 3D backbone, show, that there are the change of molecular
interaction Figure 4.
229
Figure 4. The molecular interaction of one amino acid at Z direction increases according to the order.
In doing so, the elements of (x, y, z, q) may be considered as the probabilities (pij) with
which the amino acid I presents a truncated electrostatic interaction of energy E ij , with
the amino acid j placed at a distance d ij.
(1)
The truncated electrostatic interactions are those, which occur between amino acids at a
cut-off distance (din) shorter than the half of the sum of their van der Waals radius. Here, a
shifting function (ij) was used to indicate the presence (ij = 1) or absence (ij = 0) of the
truncated electrostatic interaction between aminoacids I and j occurs by the above
interactions (Fick, 2004; Molina and Uriarte, 2004; Nordhaus and Shellenberger, 2007;
230
Philander, 2009). The theory of Doom guard Factotum states that Entropy leads everything
in the multiversity to its ultimate doom. And Entropy always wins. It can be delayed, but
never defeated. Energy consumption will not decrease, so humanity needs to find a way to
control its needs for energy, so Global Warming can be controlled if the energy
consumptions are reduced. The humanity also needs to find more efficient ways to generate
energy. These two things would help a lot towards controlling this problem, but anyway, the
increase of heat will continue. (Hasselmann, 1997) cited further that as I believe in the laws of
thermodynamics, and I don't believe in a perpetual energy source 100% efficient, the only
way to solve this problem would be to produce energy using sources outside the Earth. The
most straightforward answer for this comment is the use of Solar Energy. That is the only
actual way of reducing Global Warming on Earth currently approached. The Sun is like a
battery, there is no known way of pulling more energy than it generates, but it produces
energy enough to sustain life on Earth for billions of years yet. Using the Sun energy we
can lower the Entropy (S) of the Earth, and in this way we can reduce the problem of
increasing Global Warming of the planet. Eventually the Sun's Entropy will reach the
values so high that it will become a red giant, almost enveloping the Earth orbit, and
destroying this beautiful planet, and all traces of civilization here(Hasselmann, 1997;
Willium, 2008; Nordhaus and Shellenberger, 2007; Ptry, 1995). In the present chapter author
has described the said concept in the form of big and small delta ( and ). In India there are
sufficient program have been initiated by Prime Minister Council of Climate Change, wherein
the NAVARATNA companies are earmarked. Two percent of their profit to be invested in
the control program of Global warming as part of CSR (Corporate Social Responsibility).
Global warming can also be captured, recently by catalytic membrane process, by increasing
the culture of algae, either in algal ponds, or in photo bioreactor, various biodiesel programme
can be initiated. We can surely create a lot of power plants in our planet, which can generate
enough energy to sustain the earths life, interplanetary energy transfer methods, yet
unknown, to bring this energy from one place of earth to other, through microwave
propagation. Big Bang could also be used as a clean energy source in plasma fusion. There
are many possibilities, but currently the only feasible one, is to rely on biodiesel, and the use
of solar radiation as a source of energy. Photovolticells, Fuel cells and solar cookers are
falling in these categories. While most of the energy generated on earth comes from earthly
sources, there will still be heat being generated and the Entropy will still be increased. This
is an Entropy factor to describe Global Warming. It can be delayed but never defeated. So,
lets embrace Global Warming, because we cannot run away from it, yet. Solar power is
somewhat a reality, that human being use it to sustain humanity and mankind on the earth.
There is a long and winding road ahead of us before humanity can use the Sun as its most
important source of energy again. Decay, Death and Destruction will be always in our
doorsteps, one way or another, so being used to it. So it might be concluded, that to prevent
flood, to prevent ice melt and to prevent Global warming mankind can really develop,
several possibility and among them the use of solar and wind energy and the reductions of
population, recycling and NCER (Nonconventional Energy Resources) would be essential
and as an utmost factor (Philander, 2009).
231
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Analysis Division, FAO, 1.45-1.50. (1995).
Philander, S. G.: Encyclopedia of Global Warming and Climate Change. Nature, page 59
(2009).
Willium, M. J.: Entropy shows that Global warming should cause increase viability in the
weather, Global warming, American Physical Society, Vol.17: (2008).
Walter, K.M., S.A. Zimov, J.P. Chanton, D. Verbyla and F.S. Chapin: "Methane bubbling
from Siberian thaw lakes as a positive feedback to climate warming". Nature., 443(7107),
71-75 (2006).
Chapter 12
234
INTRODUCTION
The Global Positioning Systems (GPS) developed by US for air-land- and marinenavigation, which primarily available for US DoD (Department of Defence) and latter on it is
made available for civilian users. Nowadays most of the navigation services uses GPS tool for
movement. However, the most important element of GNSS services is safety measure which
does get affected when GPS signal passes through the ionospheric irregularities. The
ionospheric irregularities lower group velocity, or speeds up the phase velocity of the
transionospheric GPS signals. The delay or acceleration of the signal produces a range error,
eventually affecting the position solution obtained from the GPS receiver. The aurora storm is
common phenomena at high latitude which creates ionospheric irregularities and eventually
degradation in GPS performance.
AURORA
The sun continuously emits charged particles in the form of solar radiation known as
solar wind whose speed ranges from a few hundred to one thousand km/sec. In addition to
solar wind thrust, the earth is exposed to the interplanetary magnetic field (IMF) of ~5-10 nT,
most of which lies in a heliospheric equatorial plane. The earths magnetic field protects the
earth from the solar wind carrying ~40 TW of solar energy flux at ~50 Re (Re is radius of the
earth) from the center of the earth. However, a fraction of the solar wind energy (~0.4 TW)
enters the magnetosphere, known as leak in (Gordon, 1998) moreover, the amount of leak in
energy depends upon the direction of IMF, if IMF is parallel to the earths magnetic field,
then a very small fraction of solar wind energy manages to get into the magnetosphere while
in the case anti parallel IMF, the maximum amount of solar wind energy flows enters the
magnetosphere. Figure 1. depicts the anti parallel IMF with earths magnetic field and Figure
1. (A) illustrate the flow of energy as indicated by solid arrow which elongates the magnetic
field lines of the earth, termed magnetotail. The field lines reconnect at the magnetotail and
trap the energy. The trapped energy in the earths magnetic field is accelerated along the
magnetic field toward the higher-latitude illustrated in Figure 1.(B) and then strikes the
atmosphere to form the aurora. The aurora is visible at night in the form of dancing of green,
yellow, or purple lights. The shape of aurora is oval (or annulus) and therefore, it is referred
to as the auroral oval. This oval, or annulus, is centered over the magnetic pole during quiet
times. The annulus grows when the magnetosphere is disturbed. The location of the auroral
oval is generally between 60 and 70 N or S. The auroral arc extends equatorward and
maximum at 67 of geomagnetic latitude at midnight illustrated in Figure 2. (A), while at
noon it extends towards pole up to 77 latitude (Feldstein, 1963).
235
Figure 1. A represents open magnetic lines. B represents the closed magnetic lines.
Figure 2. A. Represents the equatorward extension of aurora in midnight. B. Energy precipitation (E0)
due to aurora, recorded in South Pole at 16:00 UT from TIMED satellite.
Figure 2.(B) depicts an example of auroral precipitate energy at high latitude due to
auroral storm. The auroral precipitate energy at South Pole obtained from TIMED satellite at
16:00 UT time seems to be maximum (between 7 keV to 10 keV) at midnight. The large
precipitate of auroral energy affects the electron density of the ionosphere. A strong
enhancement in TEC (Total Electron Content) was recorded on March 31, 2001 due to strong
auroral precipitate energy (Foster and Vo, 2002). The enhanced TEC adds extra phase
modulation to the satellite signal during propagation through auroral oval. The extra phase
modulation to signal generates a random fluctuation of phase in the original signal. The
random fluctuation of phase or amplitude of the signal, termed as scintillation (Hey et al.,
1946) is very common in the higher latitude due to auroral storms. The phase scintillation of
GPS signal was correlated with auroral activity in the arctic (Smith et al., 2008). The strong
scintillation not only degrades the signal but may lose lock of the satellite. This problem is
severe during the solar maximum cycle (Skone and Canon, 1999) observed RMSE (Root
Mean Square Error) of positional error from 60 to 80 cm during solar maximum period. The
236
next solar maximum cycle is expected around 2013, therefore, scientists and engineers have
been alerted to model its effects.
This research will find the threshold value of the auroral precipitate energy which may
affect the GPS-based navigation at high latitude. The experiment was conducted on a Russian
cargo ship, the M. V. Emerald from Indian Bay (69.60 S, 12.43 E) to Larsemann Hill
(69.06 S, 76.03 E). The ships route was selected within the Antarctic Circle so as to observe
auroral oval.
MEASUREMENT OF AURORA
Ground-Based Measurement
When the highly energized solar particle hits the magnetosphere, the geomagnetic field of
Earth is disturbed. The aurora is produced by the disturbed magnetic field line of the earth, so
in order to take advantage of this effect, magnetometer stations were installed across the
world at high and mid latitudes. The average value of variation of Horizontal component (H)
of earth magnetic field obtained from installed magnetometer gives an idea about the
occurrence of aurora. The commonly used magnetic index to represent magnetic storm is Kp,
and it is global index obtained every three hours. When Kp > 5, it is considered to severe case
of geomagnetic storm; Kp < 4 is a geomagnetic quiet condition; and Kp = 4 is moderate case.
The distribution aurora is not based on luminosity but on the measurement particle energy
(Hardy et al., 1985). Handy studied the zones of electrons and ion flux in terms of Magnetic
Local Time (MLT) at all level of geomagnetic activity quantified by Kp. Chubb and Hicks in
1970 used Kp to study the level and mechanism of aurora; according to them, the equator
boundary of the oval expands about 1.7 equatorward per unit of Kp on the day side of the
earth and 1.3 on the night-side while it moves by 1-3 of latitude during Substorm time
(Chubb and Hicks, 1970).
Satellite-Based Measurement
Satellites with image capturing equipment (ultraviolet image sensing) are sent into the
polar orbit to capture the snapshot of the atmosphere during their passes. Several
projects of NASAs is based on studying auroral activity; the TIMED (Thermosphere,
Ionosphere, Mesosphere, Energetic, and Dynamic) mission is one of them. TIMED is aGlobal
Ultraviolet Imager (GUVI) that provides cross-track scanned images of the Earths ultraviolet
airglow and auroral emission in the Far Ultraviolet (FUV) at wavelengths 115.0 to
180.0 nm, scanning imaging spectrograph that provides horizon-to-horizon images at
five wavelength intervals (TIMED webpage http://www.timed.jhuapl.edu, 2008). It provides
information of the ionosphere and thermosphere by monitoring all three regions: daytime
mid-latitude, night-time low- to mid-latitude ionosphere and the high-latitude auroral zone,
these regions are then characterized by energy and flux of the electrons (TIMED webpage
http://www.timed.jhuapl.edu, 2008).
237
Figure 3. A. Experimental set on the ship and GPS signal are shown passing through aurora. B. The red
line indicate the ship path from Indian Bay (69.60 S, 12.43 E) to Larsemann Hill (69.06 S, 76.03 E).
238
parameter can degrade the position solution. In this study the position of the ship is estimated
in real time using constant velocity and a constant acceleration model. In marine navigation,
the precise position solution is very important. The GPS provides a position solution that does
not degrade over time. However, the case may be different when there is an ionospheric
storm. In real time kinematics, it is difficult to obtain a high precision position solution,
because the stochastic properties of the system depend on factors, such as the ships dynamic
status and physical environment (e.g., rolling and pitching of ship on a rough sea), which are
not always fixed. Therefore, efficient Kalman filtering algorithms have become an attractive
research topic. In this study, the efficient Kalman modelling is not an issue because the
Kalman filter algorithm has been used to estimate the position solution in real time. In this
study, only the ionospheric condition related to aurora is correlated with the positional error
obtained from the GPS receiver.
The positional error of the ship in real time is computed in three steps. In the first step,
the positional solution is computed by the software of receiver; in the second step, the
position solution is estimated using constant velocity and constant acceleration model and
filtering out the noises using Kalman filter. (The algorithm of the second stage is described in
the next paragraph.) In the third stage, the position error is computed by taking the difference
of the estimated positional solution from model with Kalman filter and position solution
obtained from GPS in Cartesian coordinate frame.
A constant velocity and acceleration model is used here to measure the update of
positions, because it is assumed that the velocity of ship cannot be changed within a minute.
A 9-dimensional state vector contains three components of position (x, y, z), three
components of velocity (vx, vy, vz) and three components of acceleration (ax, ay, az). The
average of velocity and acceleration for a minute is taken as a constant velocity and
acceleration of the model for the next epoch, and each epoch is recorded at 1 sec. Figure 4.
explains the real kinematic state of the ship; consider the ship is at point P at time t1 after one
minute of initial stage the ship reached point Q. The velocity and acceleration obtained for 1
min are averaged to have one constant value which would be used to compute the position of
the ship in next epoch therefore our first computation is at 1min 1 sec. When the ship reached
to point R then the model will take the average value of velocity and acceleration obtained
between points Q and R. The probable noises are filtered out through the Kalman filter
model.
Kalman Filter
The filter estimates the process state at some time and then obtains feedback in the form
of (noisy) measurements. The mathematical process of Kalman filter falls into two groups:
Time Update equations and Measurement Update equations. The time update equations are
responsible for projecting forward solution by using the current state and error covariance
estimates to obtain a priori estimates for the next time. The second is the measurement update
equations, and it is responsible for the feedback (i.e. for incorporating a new measurement
into a priori estimate to obtain an improved a posteriori estimate). Figure 5. shows the
algorithm of the Kalman filter.
239
X k x, v x , a x , y, v y , a y , z, v z , a z
T
k
(1)
x k 1 x k tv x
t 2
ax
2
(2)
240
y k 1 y k tv y
t 2
ay
2
(3)
z k 1 z k tv z
t 2
az
2
(4)
With the help of transition matrix Fx the relation between previous and current states is
governed by
0
0
0
0
t 2
2
t 2
0
2
t 2
0
2
0
0
t 0
0 t
0 0
1 0
0 1
(5)
Wk
q xT 3
3
0
q xT 2
q xT 2
2
q xT
q yT 3
q yT
2
0
q yT 2
2
q yT
q zT 3
3
q zT 2
2
q zT 2
2
q zT
(6)
Where qx, qy, qz are the process noise variance were assumed due to troposphere, multipath
and stream of the sea (rolling and pitching).
241
242
and Schunk, 1986). The work in demonstrates the correlation of TEC derived from GPS with
TIROS precipitating energy flux over 0.320 keV (Coker et al., 1995). They mentioned that
short enhancement of TEC correlates with the precipitating electron flux. Aarons et al. (2000)
have used Polar UVI data to study the production of fluctuations in GPS signal produced by
TEC variations correlate well with increased TEC. Similar studies is seen in (Pi et al., 1997)
where author considered worldwide GPS network to investigate GPS TEC fluctuations and
present a case showing high-latitude TEC fluctuations in the evening and early morning
during a magnetic storm. The TEC fluctuation can be produced by precipitating electron flux.
The sum of this evidence is that TEC fluctuations occur in the same regions as active
auroral displays. The tracking performance of GPS receiver can be degraded by the
fluctuation of TEC and during periods of enhanced ionospheric activity in the high latitude
auroral region is very significant (Skone, 2001). Figure 6 represents the case when the GUVI
energy flux is approximately equal to 10 keV and when the GPS satellites are in the line of
sight and the degradation in position solution is greater.
The first panel of Figure 8 represents northing and easting error; the second and third
panels represent the total number satellites locked and HDOP, respectively. Results show that
when aurora activity is observed, a corresponding strong fluctuation in horizontal position is
recorded. The evidence of enhanced TEC degrades the SNR of the signal and the degraded
signal causes significant positional error. These variations are associated with smaller
intensifications in the auroral oval, which are often observed as precursors to the more intense
expansive phase velocity (Murphree et al., 1991).
Due to high auroral activity the number of satellite locked also decreases and reaches the
minimum number of 4 between 19:00 UT and 20:00 UT. With the decreases number of
satellites, the geometry is affected and increased HDOP is also a source of positional error
(Position Accuracy = HDOP x User Range Error).
The easting error in Figure 8. shows one-to-one correlation with the PRNs affected by the
auroral storm. Figure 8. also illustrates that the positional solution is smooth when there is no
auroral storm.
243
Figure 6. The upper panel represents the auroral precipitate energy and the lower represents the IPP of
the GPS satellite passing through the aurora (computed in MLT reference).
SNR (dB)
PRN:1
PRN:4
PRN:12
60
60
60
60
50
50
50
50
50
40
40
40
40
40
30
30
30
30
30
20
16
18
20
20
22 16
18
20
22
20
16
PRN:17
PRN:14
SNR (dB)
PRN:11
PRN:9
60
18
20
20
16
22
PRN:20
18
20
22
20
60
60
60
60
60
50
50
50
50
50
40
40
40
40
40
30
30
30
30
30
20
20
22 16
20
22 16
18
20
18
20
18
20
UT (hours)
20
22 16
18
Figure 7. SNR plot of all locked satellites between 18:00 22:00 UT.
20
18
20
22
PRN:28
PRN:22
22
20
16
18
20
22
DOP
ary
of S2
244
20
10
East Error
North Error
-10
00:00
02:00
04:00
06:00
08:00
10:00
14:00
16:00
18:00
20:00
22:00
00:00
14:00
16:00
18:00
20:00
22:00
00:00
14:00
16:00
18:00
20:00
22:00
00:00
10
B
5
0
00:00
02:00
04:00
06:00
08:00
10:00
12:00
HDOP
HDOP
12:00
Number of Satellite
2
0
00:00
02:00
04:00
06:00
08:00
10:00
12:00
UT(HH:MM)
Figure 8. A horizontal error in terms of Easting and Northing error B. Number of satellite locked on 28
Feb 2007, C. Horizontal Dilution of Precision (HDOP).
An estimation of position of ship is purely based on MLT confined within the auroral
plot, as indicated by the solid red arrow. The IPP of total number of visible satellite are
illustrated in the lower panel of Figure 9. During this event the maximum number of locked
satellites was 7. Figure 10. illustrates the effect of the SNR of the signal and the SNR value
obtained indicate no degradation of the signal in the absence of aurora.
The horizontal position errors are illustrated in the upper panel of Figure 11, the results
show negligible horizontal position errors in terms of easting and northing. The second panel
of the Figure 11. illustrates the number of locked satellites, and the third panel represents
HDOP to study the satellite geometry.
The results show that in the absence of auroral activity the position accuracy was fair and
good. Though position error was recorded high around 06:00 UT to 15:00 UT due to the low
number of satellites locked (4), the HDOP increased to 8.
As mentioned in the previous section, the big value of HDOP is also responsible for
positional error. Therefore, the positional error obtained is due to bad geometry.
245
Figure 9. The upper panel represents the auroral precipitate energy and the lower represents the IPP of
the GPS satellite not passing through the aurora (computed in MLT reference).
PRN:6
PRN:16
50
50
50
50
40
20
11
40
30
12
13
14
20
11
PRN:23
40
30
12
13
20
11
14
PRN:24
50
50
50
30
20
11
SNR
60
SNR
60
40
30
12
13
14
20
11
12
13
14
40
30
12
13
UT (Hours)
14
20
11
40
30
PRN:25
60
40
SNR
60
SNR
60
30
SNR
PRN:13
60
SNR
SNR
PRN:2
60
12
13
14
20
11
12
13
14
246
Horizontal Error (Meter)
-20
00:00
02:00
04:00
06:00
08:00
10:00
12:00
14:00
16:00
18:00
20:00
22:00
00:00
Number of Satellite
No. of Satellite
12
-------Data NotAvailable--------4
0
00:00
02:00
04:00
06:00
08:00
10:00
12:00
14:00
16:00
18:00
20:00
22:00
00:00
HDOP
8
HDOP
--------Data NotAvailable---------
4
2
0
00:00
02:00
04:00
06:00
08:00
10:00
12:00
14:00
16:00
18:00
20:00
22:00
00:00
UT(HH:MM)
Figure 11. A Horizontal error in terms of Easting and Northing error B. Number of satellite locked on
01 April 2007, C. Horizontal Dilution of Precision (HDOP).
247
ACKNOWLEDGMENTS
The authors wish to acknowledge the financial support from NCAOR, Goa, Ministry of
Earth Science, Govt. of India, under the Space Weather Programme at Antarctica. The
authors also acknowledge the TIMED mission for providing aurora data.
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Basu, S., S. Basu, E. MacKenzie and H. E. Whitney: Morphology of phase and intensity
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INDEX
A
abatement, 222
absorption spectroscopy, 120
access, 248
accessibility, 119
acclimatization, 131
acid, 62, 67, 97, 115, 229
active oxygen, 97, 99, 113
active site, 120
adaptability, 54
adaptation, 40, 68, 72, 74, 82, 95, 108, 139, 146,
163, 164, 169, 171
adaptations, viii, 68, 227
adenine, 119, 128
adjustment, 95, 111
advancement, viii
adverse conditions, 11, 100, 107
adverse effects, 115
adverse weather, 44
aerosols, 18, 196
Africa, 2, 3, 135, 136
age, 24, 28, 40, 41, 222, 225
agriculture, 7
Air Force, 195
air pollutants, 70
air temperature, 5, 22, 67, 69, 74, 75, 79, 141, 169,
202, 213
Alaska, 224
Alaskan North Slope, 82
alcohols, 99, 103
alfalfa, 126
algae, 11, 41, 44, 47, 49, 50, 53, 56, 58, 59, 61, 62,
63, 66, 68, 72, 74, 75, 80, 82, 83, 84, 85, 87, 95,
98, 99, 104, 109, 129, 131, 133, 164, 228, 230
algorithm, 238, 239
alien species, 41
alpha-tocopherol, 95
alters, 16, 104, 126, 146
B
bacteria, 47, 58, 68, 70, 83, 107, 118, 132, 226
bacterium, 58, 68, 70, 83
banks, 53, 54, 61, 143
barriers, 15
base, 14, 57, 97, 118, 119, 126, 203
base pair, 126
behavioral change, 223
behavioral intentions, 223
bending, 116
250
Index
C
cabbage, 143
cadmium, 123
calcium, 66
campaigns, 201
carbohydrate, 96, 97
carbohydrates, 59, 101, 121, 142, 143, 145, 148, 170
carbon, 6, 9, 10, 15, 32, 34, 39, 63, 66, 67, 77, 81,
97, 105, 108, 126, 128, 143, 146, 147, 221, 222,
224, 225, 226, 228
carbon dioxide, 6, 9, 15, 34, 105, 126, 128, 146, 147,
221, 222, 224, 226
carbon monoxide, 6
carefulness, 9
carotene, 94, 95, 98
carotenoids, 72, 82, 98, 100, 102, 131
case study, 241
cation, 120
cellulose, 100, 103, 222, 228
chaos, 221, 223, 228
chemical, 5, 9, 10, 11, 63, 222
chemical characteristics, 11
chemical properties, 222
chemical reactions, 9
251
Index
construction, 10
consumers, 11, 49, 50, 51, 54, 56, 59, 101
consumption, 68, 85, 230
consumption rates, 68, 85
contamination, 32
Continental, 32, 108, 109, 133, 142, 144, 196
contour, 209
convention, 211
convergence, 2
cooling, 15, 16, 20, 43, 55, 56, 68, 79, 176, 177, 178,
180, 183, 194, 199, 208, 221, 225, 228
cooperation, 171
coordination, 31
copper, 4
correlation, 87, 174, 180, 189, 211, 242
correlations, 247
cortex, 97, 115, 123
cosmic ray flux, 174, 175, 196
cosmic rays, 174, 195, 196
cost, 112, 223
courtship, 57
covering, 3, 44, 164
critical analysis, 11
criticism, xv
crop, 62, 72, 87, 226
crop production, 226
crops, 10, 105, 128, 226
crust, 3, 158
crystal structure, 126
crystalline, 41
crystals, 70, 167, 169, 219
Cuba, 144
culture, 230
curricula, 223
cuticle, 55, 164, 169
cycles, 40, 108, 112, 125, 142, 145, 207, 209, 213,
224
cycling, 33, 48, 58, 59, 132
cyclones, vii, 15, 16, 17, 18, 31, 39, 40, 42, 43, 46,
67, 69, 70, 79, 85
cytoplasm, 116
cytosine, 126
D
danger, 228
data set, 69
decay, 177, 189, 241, 248
decomposition, 39, 59, 67, 81, 82, 84
defence, 107, 111
deficit, 104, 200
deforestation, 224
degradation, 75, 102, 112, 234, 242, 244, 247
dehydration, 95, 143
deposition, 16
deposits, 3, 10, 123, 225
depression, 47, 141
depth, 6, 7, 26, 28, 35, 40, 42, 47, 66, 67, 73, 110,
139, 202, 203, 215
derivatives, 96, 166, 169
desiccation, 41, 55, 71, 87, 97, 99, 113, 115, 116,
127, 128, 140, 141, 142, 163, 164, 165, 169,
171, 172
destruction, 99, 100, 105
developed nations, 221, 222, 228
developing countries, 222
developing nations, 221, 228
deviation, 176, 177, 180, 183, 186, 192, 246
dew, 41, 70, 75, 77, 80, 169
DHS, 126
diffraction, 241, 248
diffusion, 28
disaster, 221
diseases, 226
displacement, 222
dissociation, 121
dissolved oxygen, 47
distribution, 10, 21, 35, 51, 55, 67, 68, 70, 84, 89, 95,
97, 99, 105, 110, 118, 121, 124, 125, 127, 132,
133, 138, 145, 149, 150, 156, 158, 159, 161,
165, 171, 187, 189, 236
diversity, 8, 39, 40, 47, 77, 80, 82, 96, 108, 147, 150,
172
DNA, 72, 107, 115, 116, 117, 118, 119, 120, 121,
122, 123, 124, 125, 126, 127, 128, 129
DNA damage, 115, 116, 117, 118, 121, 122, 126,
129
DNA lesions, 117, 118, 119, 122, 128
DNA repair, 118, 121, 125
DOI, 34, 35, 36
dominance, 62, 164, 171
drainage, 163, 165, 166, 178, 181, 187, 195
dream, 107
drinking water, 4, 222
drought, 37, 75, 77, 95, 97, 115, 141, 170, 226
dry matter, 170
drying, 47, 55, 98, 116, 128, 142
DS-1, 83
duality, 112
E
ecology, 41, 74, 79, 84, 126, 133, 144, 145, 147,
161, 166, 172
ecosystem, viii, 5, 34, 39, 44, 47, 48, 54, 58, 59, 62,
63, 67, 69, 70, 72, 73, 74, 75, 79, 81, 83, 84, 85,
122, 123, 143, 146, 226
editors, iv, xv
252
Index
effluent, 231
egg, 57
elasticity modulus, 163, 165, 168, 169, 170, 171
electric current, 192, 193
electric field, 173, 174, 180, 181, 184, 193, 194, 195
electricity, 196
electron, 10, 99, 119, 120, 123, 235, 241, 242, 248
electrons, 236
elongation, 100, 105
emission, 221, 222, 228, 236
enemies, 57
energy, 5, 8, 9, 14, 15, 16, 17, 18, 29, 39, 41, 45, 47,
48, 49, 58, 59, 66, 67, 69, 71, 72, 75, 78, 79, 80,
91, 96, 97, 98, 118, 120, 121, 132, 164, 174,
199, 200, 211, 212, 219, 221, 222, 223, 228,
229, 230, 233, 234, 235, 236, 237, 241, 242,
243, 245, 247
energy consumption, 230
energy transfer, 230
engineering, 222, 223, 226
entropy, 221, 223, 224, 226, 228, 231
environment, iv, vii, viii, ix, 1, 3, 4, 5, 7, 8, 10, 11,
13, 15, 17, 20, 22, 23, 32, 40, 41, 43, 45, 47, 48,
49, 50, 61, 67, 68, 70, 75, 77, 79, 81, 84, 85, 98,
116, 124, 126, 127, 132, 133, 146, 150, 170,
171, 223, 224, 226
environmental aspects, 224
environmental change, viii, 123, 144
environmental conditions, viii, ix, 5, 11, 91, 95, 96,
97, 99, 132, 133, 138, 139
environmental control, 143
environmental factors, 47, 96, 99, 223
environmental influences, 41
Environmental Protection Agency (EPA), 227
environmental stress, 113
environmental stresses, 113
environmentalism, 231
enzyme, 39, 115, 118, 119, 120, 127, 141
enzymes, 100, 103, 104, 115, 118, 119, 121, 140
epidermis, 114
EPR, 120
equilibrium, vii, 16
equipment, 10, 236
ERA, 35
erosion, 30
eukaryotic, 119
Europe, 2, 159
evaporation, 15, 42, 67, 71, 72, 98, 108, 150, 225
evidence, 5, 77, 98, 100, 112, 113, 176, 196, 241,
242
evolution, 32, 48, 68, 81, 85, 86, 104, 118, 122, 127,
139, 143, 223
excision, 128
F
FAD, 119, 120
families, xv, 55
fat, 57
fatty acids, 142
fauna, 39, 48, 50, 51, 63, 66, 80, 84, 85, 86, 87, 131
fertility, 66
fertilization, 75, 77, 80
fertilizers, 10
films, 223
filters, 101, 122, 124
filtration, 49
financial, 81, 171, 247
financial support, 247
fish, 57
fixation, 143
flavonoids, 96, 99, 100, 103, 113, 122, 124, 125,
128, 145, 147
flexibility, 104
flight, 58, 70
flights, 70
floods, 226
flora, viii, 3, 4, 11, 39, 40, 41, 44, 47, 48, 49, 50, 51,
56, 67, 68, 70, 71, 72, 75, 77, 79, 80, 81, 86, 87,
90, 91, 95, 96, 100, 102, 104, 105, 108, 121,
128, 131, 132, 133, 139, 143, 146, 149, 153,
157, 159, 160, 161, 227, 228
flora and fauna, 3, 4, 11, 39, 40, 41, 44, 47, 67, 68,
70, 71, 72, 79, 81, 90, 100, 227, 228
flowers, 11
fluctuations, 112, 141, 173, 189, 195, 215, 242, 247
fluorescence, 98, 112, 124
food, 11, 40, 47, 48, 50, 51, 54, 58, 59, 75, 80, 81,
101, 223, 225, 228
food chain, 11, 48, 58, 59, 80
food web, 58, 101
force, 15, 186, 187, 200
formation, vii, 7, 10, 13, 14, 15, 16, 17, 18, 22, 24,
28, 35, 39, 41, 66, 67, 68, 69, 70, 75, 77, 79, 99,
116, 117, 120, 121, 125, 126, 132, 173, 176,
177, 180, 187, 189, 194, 200, 218, 222
formula, 186
fossils, 4
fragments, 52, 53, 54, 139
253
Index
freedom, 12
freezing, 9, 16, 18, 44, 50, 55, 56, 62, 68, 95, 98, 99,
103, 113, 125, 133, 140, 141, 142, 145, 150,
193, 222
frequency distribution, 205, 206
freshwater, 28, 47, 56, 63, 68, 86, 107, 164
frost, 74, 150, 170
fruits, 11
fuel consumption, 222
functional analysis, 104, 127
fungi, 47, 56, 58, 70, 73, 86, 95, 99, 118, 131, 132,
164
fungus, 121
fusion, 75, 79, 230
G
gametophyte, 142
gene expression, 127
genes, 124, 144, 145
genetic code, 117
genetic diversity, 133
genetic information, 118
genome, 118, 119
genus, 54, 55, 57, 58, 96, 132, 146, 149, 158, 159,
160
geography, 5, 86
geometry, 110, 237, 242, 244, 246
Georgia, 87, 126, 132, 134, 136, 149, 150, 157, 159,
160, 161, 172
Germany, 90, 125, 161
germination, 71, 112
global climate change, 121
global scale, 82
global warming, 5, 7, 17, 22, 27, 33, 35, 36, 82, 219,
221, 222, 223, 224, 225, 226, 227, 228, 231
glucose, 228
glycerin, 99
glycerol, 55
google, 224
GPS, 233, 234, 235, 236, 237, 238, 241, 242, 243,
245, 247, 248
gracilis, 51, 155
graph, 205, 209
grass, 11, 99, 103, 125, 132, 143, 144
grasses, 133
gravitation, 8
gravity, 15, 16, 202
grazing, 101, 104
green alga, 61, 62, 72, 73, 80, 86, 96, 97, 99, 164
greenhouse, 28, 34, 74, 223, 224, 225, 227
greenhouse gases, 223, 224, 225
growth, 11, 14, 36, 39, 40, 44, 47, 49, 50, 51, 55, 61,
62, 63, 66, 67, 68, 73, 79, 80, 81, 83, 87, 91, 95,
97, 98, 100, 102, 104, 105, 106, 107, 108, 111,
112, 115, 121, 127, 128, 129, 140, 141, 142,
143, 146, 147, 148, 164, 177, 225
growth rate, 36
growth temperature, 98
Guangzhou, 86
H
habitat, 41, 44, 47, 50, 54, 55, 57, 61, 68, 70, 75, 78,
79, 85, 96, 97, 107, 115, 132, 138, 164, 168,
171, 225, 226
habitats, 41, 49, 51, 52, 53, 67, 95, 123, 125, 127,
131, 132, 139, 141, 142, 149, 164, 165, 172,
222
hair, 99, 144
hardness, 42, 66
harmful effects, 101, 113, 222
harvesting, 119
hazards, 9, 226, 228
health, 104, 143, 223, 226
heat loss, 200
heat transfer, 17, 78, 219
height, 6, 10, 13, 22, 23, 24, 34, 57, 67, 157, 174,
184, 185, 186, 215, 218
hemisphere, 8, 42, 57, 110, 113, 157, 160, 189, 195,
196, 197
heterogeneity, 61, 75, 79
high winds, 14, 27
histogram, 206, 207
histones, 119
history, iv, 1, 5, 11, 31, 86, 103, 127, 147, 201, 222
Holocene, 40, 86
hot springs, 99
House, 10, 12, 160, 172
human, 1, 4, 5, 9, 11, 40, 45, 57, 63, 66, 70, 82, 224,
226, 230
human activity, 1, 5, 66, 82
humidity, 9, 39, 42, 43, 47, 51, 67, 71, 79, 141, 142,
164, 169
humus, 63, 81, 95
Hungary, 127
hunting, 222
hurricanes, 225
hyaline, 136
hybrid, 223
hydrocarbons, 3
hydrogen, 99, 125
hydrogen peroxide, 99
hydroxyl, 99, 113
hydroxyl groups, 113
hypothesis, 174, 213
254
Index
I
ideal, 5, 40, 72, 73, 96, 170, 201
identification, 51, 52, 53, 87, 96, 126, 133, 161, 172
identity, 5
image, 236
images, 236, 247
imagination, 11
IMF, 174, 175, 176, 177, 178, 180, 187, 188, 189,
191, 194, 197, 234
immersion, 169
immigration, 40
Impact Assessment, 224
impulses, 193
in vitro, 125
incidence, 107, 108
income, 194
India, ix, xi, xii, xiii, xv, 1, 3, 9, 13, 18, 31, 34, 35,
39, 81, 85, 89, 90, 91, 105, 107, 131, 149, 153,
154, 155, 156, 158, 160, 163, 221, 230, 231,
233, 247
individuals, 65, 121
induction, 126, 128
industrialization, 10
industries, 226
industry, 7, 224
ingredients, 227
inhibition, 72, 84, 122
insects, 47, 54, 55, 68
institutions, xv
insulation, 75, 80, 204
integrity, 97, 100
interface, 15
interference, 1
intervention, 9
intrusions, 41
inversion, 13, 16, 20, 21, 24, 25, 29, 30, 200, 201,
213, 215, 217, 218, 219
invertebrates, 40, 50, 51, 68, 74, 75, 80, 118
ionization, 4, 174, 193, 241
iron, 3, 99, 149
irradiation, 103, 104, 105, 128, 147, 170, 193
islands, 2, 3, 39, 40, 108, 109
isolation, 41, 132, 149, 157, 160
isomers, 116, 118, 125
isotherms, 165, 170
isotope, 33
issues, iv, ix, 7, 12, 87, 231
Italy, xi, xii, 34, 86, 199
J
Jamaica, 87
Japan, 86
Jordan, 32
K
kaempferol, 113, 114
L
laboratory studies, 72
lactic acid, 58, 83
lakes, 10, 16, 39, 42, 44, 47, 49, 51, 53, 54, 56, 61,
63, 66, 78, 81, 85, 91, 98, 99, 108, 138, 139,
141, 144, 145, 164, 165, 231
landscape, 72
landscapes, 145, 146, 164
Late Pleistocene, 86
laws, 230
leaching, 145
lead, vii, 16, 17, 20, 22, 23, 24, 74, 78, 116, 132,
174, 187, 195, 225, 226
legs, 55
Lepidoptera, 85
lesions, 116, 118
lice, 56
lichen, viii, 11, 47, 49, 50, 51, 52, 53, 61, 68, 71, 74,
75, 77, 80, 82, 85, 95, 96, 101, 102, 103, 108,
115, 122, 124, 125, 131, 145, 149, 150, 153,
157, 158, 160, 163, 165, 166, 167, 168, 169,
171
life cycle, 11, 67, 68
light, 11, 27, 31, 47, 49, 50, 51, 57, 62, 66, 67, 68,
73, 74, 75, 77, 97, 98, 99, 100, 102, 106, 116,
118, 119, 120, 122, 123, 124, 142, 144, 228
lipids, 99, 100, 142
living conditions, 50
logistics, 81, 217
Louisiana, 225
low temperatures, 4, 68, 74, 83, 85, 141
luminosity, 236
lutein, 95
lying, 99, 132
lysis, 62
M
macroalgae, 102
macromolecules, 97
magnesium, 66
magnetic field, 5, 174, 175, 176, 180, 181, 195, 233,
234, 236
magnetism, 4
magnetosphere, 174, 187, 192, 193, 194, 233, 234,
236
magnitude, 176, 201
255
Index
majority, 47, 58, 99, 132, 164
mammal, 87
mammals, 48
man, 149, 176, 177
management, ix
manganese, 4
manipulation, 142
marine environment, 18
mass, vii, 14, 15, 16, 28, 34, 36, 37, 164, 178, 179,
181, 200
materials, 41, 56, 58, 66, 80, 147
matrix, 240
matter, iv, 14, 17, 22, 67, 73, 226
measurement, 5, 10, 44, 96, 143, 172, 176, 200, 202,
203, 204, 236, 238
measurements, 6, 24, 35, 36, 41, 43, 44, 72, 82, 86,
96, 127, 128, 177, 180, 185, 200, 202, 203, 238,
248
mechanical properties, 170
medulla, 115
melanin, 95, 97
melt, 4, 10, 28, 36, 45, 47, 50, 52, 97, 98, 99, 108,
116, 141, 165, 169, 201, 221, 222, 223, 225,
226, 230
melting, 7, 16, 28, 31, 36, 42, 44, 47, 74, 132, 169,
211, 221, 222, 224, 226
melts, 7, 8, 10, 14, 16, 222
membranes, 101
meta analysis, 113, 140
meta-analysis, 123, 127
metabolism, 101, 103, 104, 113, 124, 140, 143, 144
metabolites, 102, 103, 124
metals, 105
meter, 10, 62, 225, 227
microbial communities, 147
microbiota, 77, 80
microclimate, 49, 67, 125, 142
microcosms, 126, 146
microenvironments, 61
micrograms, 56
microhabitats, 81
microorganism, 34, 84
microorganisms, 11, 15, 22, 39, 47, 58, 66, 71, 75,
78, 79, 80, 83, 102, 108, 126
microscope, 50
migration, 40, 226
mineralization, 67
mission, 236, 247
mixing, 16, 22, 27, 29, 70, 85, 208, 215
MLT, 236, 237, 241, 242, 243, 244, 245
model system, 72
modelling, 32, 35, 218, 238
models, 10, 23, 34, 36, 68, 174, 199, 200, 201
N
NaCl, 167
natural disaster, 225
natural disasters, 225
natural gas, 3
natural resources, 223
navigation system, 247
negative effects, 98, 115, 121, 140
nematode, 84
Netherlands, 123, 144, 146
neutral, 233
New Zealand, 3, 90, 102, 110, 125, 144
nitrogen, 33, 61, 66, 68, 95
nitrogen fixation, 61, 68
nitrous oxide, 224, 226
Nitzschia, 92, 94
non-enzymatic antioxidants, 100
North America, 2
Norway, 82
NPL, 31, 33, 43, 81
nucleation, 36, 147
nuclei, 208
nucleic acid, 91, 100
nucleus, 118
nutrient, 40, 47, 48, 50, 59, 66, 73, 81, 91, 132
nutrients, 46, 47, 48, 50, 63, 66, 104, 112, 128
nutrition, 47
O
obstacles, 3, 74
oceans, vii, 3, 5, 10, 15, 107, 225
oil, 3, 4
operations, 217
opportunities, vii, 139
256
Index
P
P. sulcata, 155
Pacific, 2, 5, 173, 187, 189, 195, 197, 218
Paraguay, 135
parallel, 234
parasites, 56
parthenogenesis, 55, 56
pasture, 83
peat, 40
percentile, 227
periodicity, 208
permeability, 100
peroxidation, 100
petroleum, 222
phenolic compounds, 103, 113
phenylalanine, 115, 141
phosphate, 66
phospholipids, 97
phosphorus, 95
photosynthesis, 11, 50, 59, 63, 66, 68, 71, 72, 73, 75,
77, 83, 84, 85, 86, 96, 98, 100, 101, 102, 104,
105, 107, 111, 112, 121, 127, 129, 141, 142,
143, 144, 146, 164, 169, 171, 172
photosynthesize, 75, 77, 169
phycocyanin, 95, 99
phycoerythrin, 99
physical environment, 238
physical properties, 1, 4, 68
physical structure, 17
physicochemical characteristics, 63
physics, 231
Physiological, 106, 122
physiology, 87, 99, 101, 146, 163
phytoplankton, 72, 84, 101, 105
257
Index
protection, 72, 74, 75, 80, 95, 97, 98, 105, 110, 113,
115, 116, 117, 131, 132, 140, 142
protective mechanisms, 107
protective role, 97, 125
proteins, 72, 91, 100, 118, 231
protoplasm, 71
publishing, ix, xv
pure water, 4, 222
pyrimidine, 116, 119, 120, 126
Q
quality control, 212
quartz, 49
quasi-equilibrium, 180, 194
quercetin, 113, 114
R
race, 8, 223
radar, 10, 201, 248
Radiation, v, 16, 71, 72, 83, 85, 104, 105, 107, 123,
144, 203, 212
radiation damage, 99, 128
radical formation, 123
radicals, 99
radio, 1, 4, 233, 237, 248
radius, 14, 186, 229, 234
rainfall, 7, 17, 77, 80, 141, 226
reaction center, 102
reactions, 66, 223, 228
reactive oxygen, 111
reactivity, 9
real time, 238
reality, 174, 230
receptors, 72, 121
recognition, 118, 132
recommendations, iv
recovery, 105
recycling, 221, 223, 230, 231
reform, 222, 223
regenerate, 120
regions of the world, 150, 159
regression, 180, 200
regression line, 180
rehydration, 142, 143
rejection, 14, 28
religion, 223
remote sensing, 8, 33, 44, 199, 202
repair, 99, 103, 107, 111, 112, 113, 115, 116, 117,
118, 119, 120, 121, 122, 124, 126, 128, 140,
144
replication, 116, 117, 118
reproduction, 40, 81
requirements, 4, 68
researchers, viii, 6, 7, 90, 157
residues, 221, 222, 228
resilience, 121
resistance, 62, 97, 125, 150
resources, 75, 80, 87, 228
respiration, 75, 77, 82, 85, 123, 143, 164, 172
response, 32, 68, 85, 91, 95, 97, 98, 101, 104, 105,
106, 113, 122, 125, 126, 127, 129, 132, 139,
140, 146, 147, 148, 160, 164, 169, 172, 174,
176, 180, 183, 184, 187, 196, 203, 206, 219
restoration, 118
risk, 7, 116, 225
rods, 149
room temperature, 167
root, 11, 77
root system, 11
roots, 164
rotifers, 47, 50, 51, 59
roughness, 202
runoff, 225
rural development, 231
Russia, xi, xii, xiii, 7, 39, 173, 224
S
safety, 234, 247
salinity, 222
salt concentration, 170
saturation, 75, 77, 80, 169
scale system, 173, 178, 194
scarcity, 169
scatter, 175
scavengers, 97
school, 223
science, viii, 3, 4, 5, 17, 27, 32, 35
scientific observation, 89
scope, 139
sea level, 7, 8, 40, 43, 149, 173, 177, 189, 195, 197,
222, 225, 226, 227, 231
seasonal changes, 142
seasonal flu, 14
secondary metabolism, 100
secrete, 97, 99
security, 132
sediment, 58, 66
sediments, 44, 48, 51, 58, 63, 86
seed, 112, 138
seedlings, 103, 123, 124, 128, 145
selectivity, 119
sensing, 44, 82, 236, 248
sensitivity, 32, 101, 102, 126
sensors, 202
seta, 53
258
Index
structure, 4, 7, 24, 34, 40, 48, 77, 84, 88, 101, 112,
114, 121, 145, 179, 202, 203, 215, 216, 217,
219, 222, 248
style, 226
substitution, 112, 178
substrate, 39, 75, 77, 81, 121, 127
substrates, 49, 50, 66, 149
succession, 50, 73
Sun, 7, 16, 18, 22, 108, 192, 195, 230, 233
supplementation, 112
suppression, 100
surface area, 90
surface energy, 37, 203, 218, 219
surface layer, 16, 61, 173, 178, 194, 219
survival, 16, 25, 40, 41, 45, 55, 67, 68, 71, 80, 81,
107, 112, 113, 118, 131, 139, 140, 171, 222,
226
survival rate, 71
suspense, 2
sustainability, 221
Sweden, 87, 90
symbiosis, 96
synthesis, 67, 115, 140
T
target, 116, 142
taxa, 138, 149, 150, 151, 152, 153, 157, 158, 166
TBP, 117
teams, 31
techniques, 44, 248
technologies, 9
technology, viii, ix, 223
technology transfer, ix
temperature, 3, 5, 7, 9, 10, 11, 15, 16, 17, 24, 31, 34,
39, 41, 43, 45, 50, 51, 58, 59, 67, 68, 69, 74, 79,
84, 85, 86, 91, 97, 102, 106, 107, 108, 112, 122,
131, 132, 141, 142, 144, 146, 147, 148, 170,
174, 178, 180, 181, 182, 183, 184, 196, 197,
200, 201, 203, 204, 206, 208, 209, 210, 211,
213, 216, 217, 218, 219, 222, 224, 226
temperature dependence, 68
terrestrial ecosystems, 73, 101, 104, 122, 127, 143,
147
territorial, 58
territory, 57
testing, 97, 216, 218
TGA, 124
thermal energy, 228
thermodynamics, 230
thinning, 89
thymine, 124, 125
time periods, 113, 242
time series, 68, 102, 213
259
Index
tin, 4
tissue, 69, 117, 168, 169, 170
total energy, 174
toxic effect, 100
trajectory, 208
transcription, 116, 117, 119
transcription factors, 117, 119
transformation, 237
transgression, 40
translation, 84
transmission, 4, 116, 118
transparency, 43
transpiration, 67, 70
transport, 7, 14, 28, 34, 36, 78, 99, 164, 200
transportation, 3, 4, 22, 70
treatment, 101
triggers, 17, 118
tropical storms, 224
tryptophan, 120, 125
tundra, 82
turbulence, 20, 24, 66, 75, 85, 201, 202, 211
turbulent mixing, 17, 46
turgor, 116, 163, 165, 167, 168, 169, 170, 172
U
United Kingdom, xii, 33, 82, 85, 104, 218, 233
ultrastructure, 100
uniform, 108
unique features, vii, 13
United, 84, 224, 225
United Nations, 84, 224, 225
universe, 2, 7
uranium, 4
USA, 101, 102, 103, 105, 128, 202, 203, 231
USSR, 144
Ultra-Violet (UV), v, viii, ix, 6, 40, 43, 44, 46, 47,
64, 67, 72, 73, 79, 81, 84, 87, 89, 91, 95, 97, 98,
99, 100, 101, 102, 103, 104, 105, 107, 110, 111,
112, 113, 115, 116, 117, 118, 119, 121, 122,
123, 124, 125, 126, 127, 128, 131, 139, 140,
143, 144, 145, 146, 147, 231, 248
UV irradiation, 139
UV light, 43, 122
UV radiation, 6, 47, 72, 91, 96, 97, 98, 100, 104,
107, 115, 119, 122, 125, 128, 140
UV-radiation, 96, 101
V
valence, 118, 128
Valencia, 83
validation, 32, 200
W
Washington, 145, 195, 227, 231, 248
waste, 223
water ecosystems, 40
water quality, 34, 84
water resources, 223
water shortages, 225
water vapor, vii, 15, 16, 17, 224, 226
wavelengths, 8, 91, 110, 113, 116, 236
wealth, 3
weather patterns, 222
web, 135, 237
websites, 27
Western Australia, 37
Western Europe, 225
whales, 11, 57
wild type, 117
wildlife, 225
wind speeds, 24, 131, 202
windstorms, 7
workers, 55, 58, 62, 68, 70, 74
worldwide, 223, 224, 242, 248
X
xylem, 164
Y
Yale University, 82
yeast, 47, 58
yield, 112, 121
Z
zinc, 4