The diversity of life

The biosphere supports between 3 and 30 million species of plants,

animals, fungi, single-celled prokaryotes such as bacteria, and single-
celled eukaryotes such as protozoans (Figure 1). Of this total, only about
1.4 million species have been named so far, and fewer than 1 percent
have been studied for their ecological relationships and their role in
ecosystems. A little more than half the named species are insects, which
dominate terrestrial and freshwater communities worldwide; the
laboratories of systematists are filled with insect species yet to be named
and described. Hence, the relationships of organisms to
their environments and the roles that species play in the biosphere are
only beginning to be understood.

Earth's known living species

Figure 1: Estimated number of known living species. The majority of species are still unknown—i.e., yet to be
described by taxonomists.

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The organization of the biosphere

Natural groupings

This tremendous diversity of life is organized into natural ecological

groupings. As life has evolved, populations of organisms have become
separated into different species that are reproductively isolated from
one another. These species are organized through their
interrelationships into complex biological communities. The interactions
in these communities affect, and are affected by, the physical
environments in which they occur, thereby forming ecosystems through
which the energy and nutrients necessary for life flow and cycle. The mix
of species and physical environments vary across the globe, creating
ecological communities, or biomes, such as the boreal forests of North
America and Eurasia and the rainforests of the tropics. The sum total of
the richness of these biomes is the biosphere.

Processes of evolution

This hierarchical organization of life has come about through the major
processes of evolution—natural selection (the differential success of
the reproduction of hereditary variations resulting from the interaction
of organisms with their environment), gene flow (the movement of
genes among different populations of a species), and random genetic
drift (the genetic change that occurs in small populations owing to
chance). Natural selection operates on the expressed characteristics of
genetic variants found within populations, winnowing members of
the population who are less well suited to their environment from those
better suited to it. In this manner, populations become adapted to their
local ecosystems, which include both the physical environment and the
other species with which they interact in order to survive and reproduce.

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The genetic variation that is necessary for a species to adapt to the
physical environment and to other organisms arises from new mutations
within populations, the recombination of genes during sexual
reproduction, and the migration of and interbreeding with individuals
from other populations. In very small populations, however, some of that
variation is lost by chance alone through random genetic drift. The
combined result of these evolutionary processes is that after many
generations populations of the same species have widely divergent
characteristics. Some of these populations eventually become so
genetically different that their members cannot successfully interbreed,
resulting in the evolution of a separate species (speciation).

The diversification of life through local adaptation of populations and

speciation has created the tremendous biodiversity found on Earth. In
most regions 1 square kilometre (0.4 square mile)
will harbour hundreds—in some places even thousands—of species. The
interactions between these species create intricate webs of relationships
as the organisms reciprocally evolve, adapting to one another and
becoming specialized for their interactions. Natural communities of
species reflect the sum of these species’ interactions and the ongoing
complex selection pressures they constantly endure that drive their
evolution. The many ecological and evolutionary processes that affect
the relationships among species and their environments
render ecology one of the most intricate of the sciences. The answers to
the major questions in ecology require an understanding of the relative
effects of many variables acting simultaneously.

The importance of the biosphere

The continued functioning of the biosphere is dependent not only on the

maintenance of the intimate interactions among the myriad species
within local communities but also on the looser yet crucial interactions
of all species and communities around the globe. Earth is blanketed with
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so many species and so many different kinds of biological communities
because populations have been able to adapt to almost any kind
of environment on Earth through natural selection. Life-forms have
evolved that are able to survive in the ocean depths, the frigid conditions
of Antarctica, and the near-boiling temperatures of geysers. The great
richness of adaptations found among different populations and species
of living organisms is Earth’s greatest resource. It is a richness that has
evolved over millions of years and is irreplaceable.

It is therefore startling to realize that our inventory of Earth’s diversity is

still so incomplete that the total number of living species cannot be
estimated more closely than between 3 and 30 million species. Decades
of continuous research must be carried out by systematists, ecologists,
and geneticists before the inventory of biodiversity provides a more
accurate count. The research has been slow. Only recently, as
the extinction rate of species has been increasing rapidly, have societies
begun to realize the interdependence of species. To sustain life on Earth,
more than the few animal and plant species used by humans must be
preserved. The flow of energy and the cycling of nutrients through
ecosystems, the regulation of populations, and the stability of biological
communities, all of which support the continued maintenance of life,
rely on the diversity of species, their adaptations to local physical
conditions, and their coevolved relationships.

Despite the limited scientific knowledge of most species, ecological

studies during the 20th century made great headway in unraveling the
mechanisms by which organisms coevolve with one another and adapt
to their physical environment, thereby shaping the biosphere. Each new
decade has produced a steady stream of studies showing that the
biological and physical elements of Earth are more interconnected than
had been previously thought. Those studies also have shown that often
the most seemingly insignificant species are crucial to the stability of
communities and ecosystems. Many seemingly obscure species are at
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risk worldwide of being dismissed as unimportant. The effect that the
loss of species will have on ecosystems is appreciated only by
understanding the relationships between organisms and
their environments and by studying the ecological and evolutionary
processes operating within ecosystems.

The need to understand how the biosphere functions has never been
greater. When human population levels were low and technological
abilities crude, societies’ impact on the biosphere was relatively small.
The increase in human population levels and the harvesting of more of
Earth’s natural resources has altered this situation, especially in recent
decades. Human activities are causing major alterations to the patterns
of energy flow and nutrient cycling through ecosystems, and these
activities are eliminating populations and species that have not even
been described but which might have been of central importance to the
maintenance of ecosystems.

The biologist Edward O. Wilson, who coined the term biodiversity,

estimated conservatively that in the late 20th century at least 27,000
species were becoming extinct each year. The majority of these were
small tropical organisms. The impact that this freshet of extinctions
would have on the biosphere is akin to receiving a box of engine parts
and discarding a portion of them before reading the directions, assuming
that their absence will have no negative repercussions on the running of
the engine. The following sections describe how many of the biological
and physical parts fit together to make the engine of the biosphere run
and why many seemingly obscure species are important to the long-term
functioning of the biosphere.

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Resources of the biosphere

The flow of energy

The photosynthetic process

Life on Earth depends on the harnessing of solar energy by the process

of photosynthesis. Photosynthetic plants convert solar energy into
the chemical energy of living tissue, and that stored chemical energy
flows into herbivores, predators, parasites, decomposers, and all other
forms of life (see also photosynthesis). In the photosynthetic process,
light energy is absorbed by the chlorophyll molecules of plants to
convert carbon dioxide and water into carbohydrates and oxygen gas.
Proteins, fats, nucleic acids, and other compounds also are synthesized
during the process, as long as elements such as nitrogen, sulfur, and
phosphorus are available.

Efficiency of solar energy utilization

Most solar energy occurs at wavelengths unsuitable for photosynthesis.

Between 98 and 99 percent of solar energy reaching Earth is reflected
from leaves and other surfaces and absorbed by other molecules, which
convert it to heat. Thus, only 1 to 2 percent is available to be captured
by plants. The rate at which plants photosynthesize depends on the
amount of light reaching the leaves, the temperature of
the environment, and the availability of water and other nutrients such
as nitrogen and phosphorus. The measurement of the rate at which
organisms convert light energy (or inorganic chemical energy) to the
chemical energy of organic compounds is called primary productivity.
Hence, the total amount of energy assimilated by plants in
an ecosystem during photosynthesis (gross primary productivity) varies
among environments. (Productivity is often measured by an increase

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in biomass, a term used to refer to the weight of all living organisms in
an area. Biomass is reported in grams or metric tons.)

Much of the energy assimilated by plants through photosynthesis is not

stored as organic material but instead is used during cellular respiration.
In this process organic compounds such as carbohydrates, proteins, and
fats are broken down, or oxidized, to provide energy (in the form
of adenosine triphosphate [ATP]) for the cell’s metabolic needs. The
energy not used in this process is stored in plant tissues for further use
and is called net primary productivity. About 40 to 85 percent of gross
primary productivity is not used during respiration and becomes net
primary productivity. The highest net primary productivity in terrestrial
environments occurs in swamps and marshes and tropical rainforests;
the lowest occurs in deserts. In aquatic environments, the highest net
productivity occurs in estuaries, algal beds, and reefs. Consequently,
these environments are especially critical for the maintenance of
worldwide biological productivity.

Energy transfers and pyramids

A small amount of the energy stored in plants, between 5 and 25 percent,

passes into herbivores (plant eaters) as they feed, and a similarly small
percentage of the energy in herbivores then passes into carnivores
(animal eaters). The result is a pyramid of energy, with most energy
concentrated in the photosynthetic organisms at the bottom of food
chains and less energy at each higher trophic level (a division based on
the main nutritional source of the organism; see community ecology:
Trophic pyramids and the flow of energy). Some of the remaining energy
does not pass directly into the plant-herbivore-carnivore food chain but
instead is diverted into the detritus food chain. Bacteria, fungi,
scavengers, and carrion eaters that consume detritus (detritivores) are
all eventually consumed by other organisms.

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The rate at which these consumers convert the chemical energy of their
food into their own biomass is called secondary productivity.
The efficiency at which energy is transferred from one trophic level to
another is called ecological efficiency. On average it is estimated that
there is only a 10 percent transfer of energy (Figure 2).

ecosystem energy transfer

Figure 2: Transfer of energy through an ecosystem. At each trophic level only a small proportion of energy
(approximately 10 percent) is transferred to the next level. Encyclopædia Britannica, Inc.

Energy is lost in several ways as it flows along these pathways

of consumption. Most plant tissue is uneaten by herbivores, and
this stored energy is therefore lost to the plant-herbivore-carnivore food
chain. In terrestrial communities less than 10 percent of plant tissue is
actually consumed by herbivores. The rest falls into the detritus
pathway, although the detritivores consume only some of this decaying
tissue. Oil and coal deposits are major repositories of this unused plant
energy and have accumulated over long periods of geologic time.

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The efficiency by which animals convert the food they ingest into energy
for growth and reproduction is called assimilation
efficiency. Herbivores assimilate between 15 and 80 percent of the plant
material they ingest, depending on their physiology and the part of the
plant that they eat. For example, herbivores that eat seeds and young
vegetation high in energy have the highest assimilation efficiencies,
those that eat older leaves have intermediate efficiencies, and those that
feed on decaying wood have very low efficiencies. Carnivores generally
have higher assimilation efficiencies than herbivores, often between 60
and 90 percent, because their food is more easily digested.

The overall productivity of the biosphere is therefore limited by the rate

at which plants convert solar energy (about 1 percent) into chemical
energy and the subsequent efficiencies at which other organisms at
higher trophic levels convert that stored energy into their own biomass
(approximately 10 percent). Human-induced changes in net primary
productivity in the parts of the biosphere that have the highest
productivity, such as estuaries and tropical moist forests, are likely to
have large effects on the overall biological productivity of Earth.

Nutrient cycling

The cells of all organisms are made up primarily of six major elements
that occur in similar proportions in all life-forms. These elements—
hydrogen, oxygen, carbon, nitrogen, phosphorus, and sulfur—form the
core protoplasm of organisms, and the first four of these elements make
up about 99 percent of the mass of most cells. Additional elements,
however, are also essential to the growth of organisms. Calcium and
other elements help to form cellular support structures such as shells,
internal or external skeletons, and cell walls. Chlorophyll molecules,
which allow photosynthetic plants to convert solar energy into chemical
energy, are chains of carbon, hydrogen, and oxygen compounds built
around a magnesium ion. Altogether, 16 elements are found in all
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organisms; another eight elements are found in some organisms but not
in others.

These bioelements combine with one another to form a wide variety of

chemical compounds. They occur in organisms in higher proportions
than they do in the environment because organisms capture them,
concentrating and combining them in various ways in their cells, and
release them during metabolism and death. As a result, these essential
nutrients alternate between inorganic and organic states as they rotate
through their respective biogeochemical cycles. These cycles can include
all or part of the following: the atmosphere, which is made up largely of
gases including water vapour; the lithosphere,
which encompasses the soil and the entire solid crust of Earth; and
the hydrosphere, which includes lakes, rivers, and oceans.
A portion of the elements are bound up in limestone and in the minerals
of other rocks and are unavailable to organisms. The slow processes
of weathering and erosion eventually release these elements to enter
the cycle. For most of the major nutrients, however, organisms not only
intercept the elements moving through the biosphere, but they actually
drive the biogeochemical cycles (Figure 3).

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energy flow
Figure 3: General paths of energy flow in the biosphere.
Encyclopædia Britannica, Inc.

The movement of nutrients through the biosphere is different from the

transfer of energy because, whereas energy flows through the biosphere
and cannot be reused, elements are recycled. The same atoms of carbon
or nitrogen may, over the course of eons, move repeatedly between
organisms, the atmosphere, the soil, and the oceans. Carbon released
as carbon dioxide by an animal may remain in the atmosphere for 5 or
10 years before being taken up by another organism, or it may cycle
almost immediately back into a neighbouring plant and be used
during photosynthesis.

The carbon cycle

Life is built on the conversion of carbon dioxide into the carbon-based

organic compounds of living organisms. The carbon cycle illustrates the
central importance of carbon in the biosphere. Different paths of the
carbon cycle recycle the element at varying rates. The slowest part of the

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cycle involves carbon that resides in sedimentary rocks, where most
of Earth’s carbon is stored. When in contact with water that is acidic (pH
is low), carbon will dissolve from bedrock; under neutral conditions,
carbon will precipitate out as sediment such as calcium carbonate
(limestone). This cycling between solution and precipitation is the
background against which more rapid parts of the cycle occur.

Short-term cycling of carbon occurs in the continual physical exchange

of carbon dioxide (CO2) between the atmosphere and hydrosphere.
Carbon dioxide from the atmosphere becomes dissolved in water (H2O),
with which it reacts to form carbonic acid (H2CO3), which dissociates into
hydrogen ions (H+) and bicarbonate ions (HCO3−), which further
dissociate into hydrogen and carbonate ions (CO32−). The more alkaline
the water (pH above 7.0 is alkaline), the more carbon is present in the
form of carbonate, as is shown in the following reversible reactions:

At the same time, carbon dioxide in the water is continually lost to the
atmosphere. The exchange of carbon between the atmosphere and
hydrosphere links the remaining parts of the cycle, which are the
exchanges that occur between the atmosphere and terrestrial organisms
and between water and aquatic organisms.

The biological cycling of carbon begins as photosynthetic

organisms assimilate carbon dioxide or carbonates from the
surrounding environment. In terrestrial communities, plants convert
atmospheric carbon dioxide to carbon-based compounds
through photosynthesis (see above The photosynthetic process). During
this process, plants cleave the carbon from the two oxygen molecules
and release the oxygen back into the surrounding environment. Plants

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are thus primarily responsible for the presence of atmospheric oxygen.
In aquatic communities, plants use dissolved carbon in the form of
carbonates or carbon dioxide as the source of carbon for photosynthesis.
Once carbon has been assimilated by photosynthetic organisms, as well
as by the animals that eat them, it is released again in the form of carbon
dioxide as these organisms respire. The release of carbon dioxide into
the atmosphere or hydrosphere completes the biological part of the
carbon cycle.

The pathways of the global carbon cycle, however, are never completely
balanced. That is to say, carbon does not move in and out of all parts of
the biosphere at equal rates. Consequently, over time some parts of the
biosphere accumulate more carbon than others, thereby serving as
major accessible carbon reservoirs. In preindustrial times the major
reservoirs of carbon were the deep and shallow portions of the ocean;
the soil, detritus, and biota of the land; and the atmosphere. The oceans
were, and still are, the greatest reservoirs of carbon. Because marine
phytoplankton have such short life cycles, the carbon in the ocean cycles
rapidly between inorganic and organic states.

In terrestrial environments, forests are the largest carbon reservoirs. Up

to 80 percent of the aboveground carbon in terrestrial communities and
about a third of belowground carbon are contained within forests. Unlike
the oceans, much of this carbon is stored directly in the tissues of plants.
High-latitude forests include large amounts of carbon not only in
aboveground vegetation but also in peat deposits. Forests at high and
low latitudes particularly are important reservoirs of carbon. An
estimated one-half of the carbon in forests occurs in high-latitude
forests, and a little more than one-third occurs in low-latitude forests.
The two largest forest reservoirs of carbon are the vast expanses
in Russia, which hold roughly 25 percent of the world’s forest carbon,
and the Amazon basin, which contains about 20 percent.

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Until recent centuries, the equilibrium between the carbon in the
world’s forests and in the atmosphere remained constant. Samples of
carbon dioxide trapped in ice during the past 1,000 years and direct
measurements of carbon dioxide in the atmosphere had remained fairly
constant until the 18th century. However, the cutting of much of the
world’s forest is, along with the increase in consumption of fossil fuels
attendant on the Industrial Revolution, has resulted in a disruption of the
balance between the amount of carbon dioxide in the forests and in the
atmosphere. The concentration of atmospheric carbon dioxide has been
increasing steadily (Figure 4); currently the rate of increase is about 4
percent per decade (see global warming and climatic variation and
change). If human activities continue to alter the relative sizes of the
carbon reservoirs worldwide, they are likely to have large effects on the
carbon cycle and other biogeochemical cycles. Large-
scale deforestation in Russia and the Amazon basin are likely to have
particularly significant effects on global carbon storage and cycling.
The warming of global temperatures also is changing which ecosystems
act as long-term sinks for carbon and which act as sources for carbon
dioxide in the atmosphere. For example, the Arctic tundra, with large
amounts of carbon stored in its soils, has been a net sink for carbon
dioxide during long periods of geologic time. The recent warming of
many Arctic regions, however, has accelerated the rate of soil
decomposition, transforming these Arctic areas into potential sources of
atmospheric carbon dioxide.

The complex cycle of carbon and the varying sizes of carbon reservoirs
illustrate some of the reasons it has been so difficult to predict the
effects that increased atmospheric carbon will have on global change.

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The nitrogen cycle

Nitrogen is one of the elements most likely to be limiting to plant growth.

Like carbon, nitrogen has its own biogeochemical cycle, circulating
through the atmosphere, lithosphere, and hydrosphere (Figure 5).
Unlike carbon, which is stored primarily in sedimentary rock, most
nitrogen occurs in the atmosphere as an inorganic compound (N2). It is
the predominant atmospheric gas, making up about 79 percent of the
volume of the atmosphere. Plants, however, cannot use nitrogen in its
gaseous form and are able to assimilate it only after it has been
converted to ammonia (NH3) and nitrates (NO3−). This reductive process,
called nitrogen fixation, is a chemical reaction in which electrons are
picked up from another molecule. A small amount of nitrogen is fixed by
lightning, but most of the nitrogen harvested from the atmosphere is
removed by nitrogen-fixing bacteria and cyanobacteria (formerly
called blue-green algae).

nitrogen cycle
The nitrogen cycle transforms diatomic nitrogen gas into ammonium, nitrate, and nitrite compounds. Encyclopædia
Britannica, Inc.

15 | P a g e
Certain species of nitrogen-fixing bacteria can coexist intimately
(symbiotically) with legumes and other plants, providing the plants with
necessary nitrogen (Figure 6). In this symbiotic association,
the bacteria become encased in nodules that grow on the roots of
plants, through which nitrogen that has been fixed by the resident
bacteria is obtained. Cyanobacteria have developed similar relationships
with various life-forms, such as liverworts, hornworts, cycads, and at
least one genus of flowering plants (Gunnera). Their symbiotic
relationship with fungi has earned its own designation—the coexistent
species are called lichen.

nitrogen fixation
Figure 6: (Right) The roots of an Austrian winter pea plant (Pisum sativum) with nodules harbouring nitrogen-fixing
bacteria (Rhizobium). (Left) Root nodules develop as a result of a symbiotic relationship between rhizobial bacteria
and the root hairs of the plant. (A) The bacteria recognize the root hairs and begin to divide, (B) entering the root
through an infection thread that allows bacteria to enter root cells, (C) which divide to form the nodule.
(Left) Encyclopædia Britannica, Inc.; (right) photograph, © John Kaprielian, The National Audubon Society
Collection/Photo Researchers

16 | P a g e
Other microorganisms perform important tasks that propel the nitrogen
cycle along. Although plants can assimilate ammonia as well as nitrates,
most of the ammonia in the soil is converted to nitrites (NO2−) and then
to nitrates by certain aerobic bacteria through the oxidative process
of nitrification. Once nitrogen has been assimilated by plants, it can be
converted to organic forms, such as amino acids and proteins. Animals
can use only organic nitrogen, which they obtain by consuming plants or
other animals. As these organisms die, certain microbes such as
detritivores are able to participate in the decomposition of organic
nitrogen into ammonia (ammonification), providing a constant supply of
ammonia to be used in the process of nitrification. Although the fixation
of atmospheric nitrogen is an essential part of the nitrogen cycle,
ammonification and nitrification are the predominant methods by which
organic nitrogen is prevented from returning to the atmosphere and is
kept cycling through the biosphere.

Some nitrogen does return to the atmosphere, however, as denitrifying

bacteria break down nitrates to obtain oxygen, thereby releasing
gaseous N2. Nitrogen is also lost from plants and soil in
terrestrial environments via other routes, including erosion, runoff,
volatilization of ammonia into the atmosphere, and leaching from soils
into lakes and streams. Eventually some of these nutrients reach the
oceans as rivers flush them onto the ocean surface.

The sulfur cycle

Sulfur is found in all living organisms as a constituent of some proteins,

vitamins, and hormones. Like carbon and nitrogen, sulfur cycles between
the atmosphere, lithosphere, and hydrosphere; but, unlike these two
other elements, it has major reservoirs in both the atmosphere and the
lithosphere. As is true in the nitrogen cycle, the activities of
microorganisms are crucial in the global cycling of this nutrient.

17 | P a g e
The process begins with geochemical and meteorologic processes such
as the weathering of rock. When sulfur is released from the rock and
comes in contact with air, it is converted into sulfate (SO4), which is taken
up by plants and microorganisms and converted into organic forms.
Animals acquire these organic forms of sulfur from their foods. When
organisms die and decompose, some of the sulfur enters the tissues of
microorganisms and some is released again as sulfate. There is, however,
a continual loss of sulfur from terrestrial ecosystems as some of it drains
into lakes and streams and eventually into the ocean as runoff.
Additional sulfur enters the ocean through fallout from the atmosphere.
Once in the ocean, some of the sulfur cycles through
marine communities as it moves through food chains, some reenters the
atmosphere, and some is lost to the ocean depths as it combines with
iron to form ferrous sulfide (FeS), which is responsible for the black
colour of marine sediments. Sulfur reenters the atmosphere naturally in
three major ways: sea spray releases large amounts of the element from
the ocean into the atmosphere; anaerobic respiration by sulfate-
reducing bacteria causes the release of hydrogen sulfide (H2S) gas
especially from marshes, tidal flats, and similar environments in which
anaerobic microorganisms thrive; and volcanic activity releases
additional but much smaller amounts of sulfur gas into the atmosphere.
Since the Industrial Revolution, human activities have contributed
significantly to the movement of sulfur from the lithosphere to the
atmosphere as the burning of fossil fuels and the processing of metals
have occasioned large emissions of sulfur dioxide. Oxides of sulfur and
nitrogen contribute to the acid rain that is common downwind from
these industrial activities.

The cycling of phosphorus and other essential nutrients

Most other major nutrients such as phosphorus, potassium, magnesium,

iron, and calcium enter terrestrial communities through the weathering
of bedrock. These nutrients lack a volatile gaseous state. Consequently,
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they cycle through the biosphere differently from carbon, nitrogen, and
sulfur, all of which sometimes occur as volatile gases. Of the nonvolatile
nutrients, phosphorus is the one that most often limits plant growth,
especially in aquatic environments.

Phosphorus and the other nonvolatile elements move unidirectionally

from land, through aquatic environments, into ocean sediments. Most
phosphorus cycling occurs between the surface and depths of the ocean.
When near the surface, phosphorus is taken up by the plankton and
passed through the food chain. It cycles back to the ocean bottom as
individuals die and fall to the ocean floor,
releasing assimilated phosphorus. Much of this element gradually
accumulates in the ocean sediment as particulate phosphorus and is
eventually brought back to the surface only through ocean upwelling and
tectonic activity. The ocean sediments are therefore by far the greatest
reservoirs of phosphorus.

In terrestrial ecosystems, much of the available phosphorus moves in a

closed cycle between living organisms and the organic debris in the
soil. Phosphate (PO43−) is the only important inorganic form involved in
this cycle. Microorganisms in the soil break down litter and other organic
matter, thereby releasing phosphate, which is then taken up by plants
and released again when they die and decompose. Soils differ in the
amount of phosphorus they contain, and in some phosphorus-poor soils
almost all the available phosphorus resides in living organisms and
organic debris. In some tropical forests, such as those in parts of Brazil,
so much of the phosphorus is contained in living organisms that the
clearing of forests eliminates most of this element. As a result, the plant
communities cannot recover, and crops cannot be grown.

The addition of phosphorus to soils poor in this nutrient and the use of
phosphorus-rich detergents have had a great impact on the phosphorus
cycle in many ecosystems. Runoff from agricultural fields and the
19 | P a g e
emptying of sewage has introduced so much extra phosphorus to rivers
and lakes that populations of plants and microorganisms have grown
explosively, sometimes creating a solid mat that extends over the surface
of the water. This growth increases the amount of organic debris in the
water, which can lead to a decrease in the available oxygen, resulting in
suffocation of fish and other animals.

The hydrologic cycle

A portion of the biogeochemical cycle of all elements involves time spent

cycling through the hydrosphere. Water itself cycles within the
biosphere. (For a detailed discussion of the hydrologic
cycle see hydrosphere: The hydrologic cycle.) Unlike the cycles of the
other major nutrients, however, the hydrologic, or water, cycle would
continue in some form even in the absence of living organisms. Most
of Earth’s water is in its core, in the sedimentary rocks near its surface,
and in the ocean. A minute percentage of the water, however,
continually cycles through the atmosphere, oceans, and terrestrial
environments mainly by the processes of evaporation and precipitation.
This part of the hydrologic cycle is driven by solar energy. Water
evaporates from both the aquatic and terrestrial environments as it is
heated by the Sun’s energy. The rates of evaporation and precipitation
depend on solar energy, as do the patterns of circulation of moisture in
the air and currents in the ocean. Evaporation exceeds precipitation over
the oceans, and this water vapour is transported by the wind over land,
where it returns to the land through precipitation. The water falling onto
terrestrial environments seeps into the ground or runs off into lakes and
streams and eventually empties into the oceans, carrying with it many of
the other major nutrients. Water also reenters the atmosphere through
the evaporative loss of water by plants (transpiration).

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Links among the cycles

Although the overall pattern of cycling of all the major elements is now
known, there is still much to learn about the relative importance of the
different stages of each cycle. For example, there is considerable debate
concerning which ecosystems act as the major sources of carbon for the
atmosphere and which act as sinks by accumulating more carbon than
they release. The ways in which the different cycles interact with one
another also must be minutely studied. It has been discovered that sulfur
availability influences the rate of nitrogen accumulation in plants and
nitrogen availability influences phosphorus uptake. All three elements
are thought to influence the rate of carbon accumulation by plants. As a
result, changes in any one of these nutrient cycles influence the other
cycles as well.

The effects that disruptions in these cycles may have within the
biosphere are not clearly understood. Natural geologic phenomena, such
as ice ages and major periods of volcanic activity, have repeatedly
disturbed these cycles throughout Earth history. Many human activities
may have impacts of similar scope. Deforestation, the burning
of fossil fuels, and increased fertilization are disturbing these cycles.
These anthropogenic disturbances have increased atmospheric levels
of carbon dioxide (Figure 4), decreased ozone (O3) levels, and
undermined the natural equilibrium of streams and lakes by excessive
nutrient enrichment from sewage, fertilizers, and factory waste (cultural
eutrophication). Gleaning more information about the biogeochemical
cycles and their interactions has become increasingly important now
that the effects of human activities are becoming more apparent.
Another potential effect that may result from human intrusions in
the environment is global warming. Carbon dioxide in the atmosphere
has the ability to act as an insulator, preventing some of Earth’s heat,
absorbed from solar radiation, from escaping back into space. This
process, known as the greenhouse effect, is suspected of
21 | P a g e
being enhanced by rising levels of atmospheric carbon dioxide (Figure 4),
which have resulted in part from the combustion of fossil fuels and the
clearing and burning of tropical forests. This increase in atmospheric
carbon dioxide and other so-called greenhouse gases could raise the
overall global temperature, causing the polar ice caps to melt, sea levels
to rise, and Earth’s precipitation to be redistributed.

The complexity and interconnectedness of each of the biogeochemical

cycles make it difficult to pinpoint how any one human activity is altering
the cycles; nevertheless, the majority of those who study these
fluctuations agree that this is happening. Disagreements generally
concern the extent to which various activities affect particular cycles and
what the long-term consequences of these disturbances will be.
John N. Thompson

Environmental conditions

Most organisms are limited to either a terrestrial or an

aquatic environment. An organism’s ability to tolerate local conditions
within its environment further restricts its distribution. One parameter,
such as temperature tolerance, may be important in determining the
limits of distribution, but often a combination of variables, such as
temperature tolerance and water requirements, is important. Extreme
environmental variables can evoke physiological
and behavioral responses from organisms. The physiological response
helps the organism maintain a constant internal environment
(homeostasis), while a behavioral response allows it to avoid the
environmental challenge—a fallback strategy if homeostasis cannot be
maintained.

The ways in which modern living organisms tolerate environmental

conditions reflect the aquatic origins of life. With few exceptions, life
cannot exist outside the temperature range at which water is a liquid.
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Thus, liquid water, and temperatures that maintain water as a liquid, are
essential for sustaining life. Within those parameters, the concentrations
of dissolved salts and other ions, the abundance of respiratory gases,
atmospheric or hydrostatic pressure, and rate of water flow all influence
the physiology, behaviour, and distribution of organisms.

Temperature

Temperature has the single most important influence on the distribution

of organisms because it determines the physical state of water. Most
organisms cannot live in conditions in which the temperature remains
below 0 °C or above 45 °C for any length of time. Adaptations have
enabled certain species to survive outside this range—
thermophilic bacteria have been found in hot springs in which the
temperatures may approach the boiling point, and certain polar mosses
and lichens can tolerate temperatures of −70 °C—but these species are
the exceptions. Few organisms can remain for long periods at
temperatures above 45 °C, because organic molecules such as proteins
will begin to denature. Nor are temperatures below
freezing conducive to life: cells will rupture if the water they contain
freezes.

Most organisms are not able to maintain a body temperature that is

significantly different from that of the environment. Sessile organisms,
such as plants and fungi, and very small organisms and animals that
cannot move great distances, therefore, must be able to withstand the
full range of temperatures sustained by their habitat. In contrast, many
mobile animals employ behavioral mechanisms to avoid extreme
conditions in the short term. Such behaviours vary from simply moving
short distances out of the Sun or an icy wind to large-scale migrations.
Some types of animals employ physiological mechanisms to maintain a
constant body temperature, and two categories are commonly
distinguished: the term cold-blooded is understood to refer to reptiles
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and invertebrates, and warm-blooded is generally applied to mammals
and birds. These terms, however, are imprecise; the more accurate
terms, ectotherm for cold-blooded and endotherm for warm-blooded,
are more useful in describing the thermal capabilities of these animals.
Ectotherms rely on external sources of heat to regulate their body
temperatures, and endotherms thermoregulate by generating heat
internally.

Terrestrial ectotherms utilize the complex temperature profile of the

terrestrial environment to derive warmth. They can absorb solar
radiation, thus raising their body temperatures above that of the
surrounding air and substrate (Figure 7), unlike the aquatic ectotherm,
whose body temperature is usually very close to that of the
environment. As this solar radiation is taken up, physiological
mechanisms contribute to the regulation of heat—peripheral blood
vessels dilate and heart rate increases. The animal also may employ
behavioral mechanisms, such as reorienting itself toward the Sun or
flattening its body and spreading its legs to maximize its surface area
exposure. At night, loss of heat may be reduced by other behavioral and
physiological mechanisms—the heart rate may slow, peripheral blood
vessels may constrict, surface area may be minimized, and shelter may
be sought.

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energy exchange
Figure 7: Energy exchange between a terrestrial reptile and the environment.
Encyclopædia Britannica, Inc.

Endotherms maintain body temperature independently of the

environment by the metabolic production of heat. They generate heat
internally and control passive heat loss by varying the quality of their
insulation or by repositioning themselves to alter their effective surface
area (i.e., curling into a tight ball). If heat loss exceeds heat
generation, metabolism increases to make up the loss. If heat generation
exceeds the rate of loss, mechanisms to increase heat loss by
evaporation occur. In either case, behavioral mechanisms can be
employed to seek a more suitable thermal environment.

To survive for a limited period in adverse conditions, endotherms may

employ a combination of behavioral and physiological mechanisms. In
cold weather, which requires an increase in energy consumption, the
animal may enter a state of torpor in which its body temperature,
metabolism, respiratory rate, and heart rate are depressed. Long-term
winter hypothermia, or hibernation, is an extended state of torpor that

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some animals use as a response to cold conditions. Torpor and
hibernation free the animals from energetically expensive maintenance
of high body temperatures, saving energy when food is limited.

Another form of torpor, estivation, is experienced by animals in response

to heat stress. This state is seen more often in ectothermic animals than
in endotherms, but in both the stimulus for estivation is usually a
combination of high temperatures and water shortage.

Humidity

Most terrestrial organisms must maintain their water content within

fairly narrow limits. Water commonly is lost to the air
through evaporation or, in plants, transpiration. Because most water
loss occurs by diffusion and the rate of diffusion is determined by the
gradient across the diffusion barrier such as the surface of a leaf or skin,
the rate of water loss will depend on the relative humidity of the air.
Relative humidity is the percent saturation of air relative to its total
saturation possible at a given temperature. When air is totally saturated,
relative humidity is said to be 100 percent. Cool air that is completely
saturated contains less water vapour than completely saturated warm
air because the water vapour capacity of warm air is greater (see climate:
Atmospheric humidity). Diffusion gradients across skin or leaves,
therefore, can be much steeper in summer when the air is warm,
rendering evaporative water loss a much more serious problem in
warm environments than in cool environments. Nevertheless, rates of
water loss are higher in dry air (conditions of low relative humidity) than
in moist air (conditions of high relative humidity), regardless of the
temperature.

Water loss from evaporation must be compensated by water uptake

from the environment. For most plants, transpirational water loss is
countered by the uptake of water from the soil via roots. For animals,
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water content can be replenished by eating or drinking or by uptake
through the integument. For organisms living in dry environments, there
are many morphological and physiological mechanisms that reduce
water loss. Desert plants, or xerophytes, typically have reduced leaf
surface areas because leaves are the major sites of transpiration. Some
xerophytes shed their leaves altogether in summer, and some are
dormant during the dry season.

Desert animals typically have skin that is relatively impervious to water.

The major site of evaporation is the respiratory exchange surface, which
must be moist to allow the gaseous exchange of oxygen and carbon
dioxide. A reduction in amount of water lost through respiration can
occur if the temperature of the exhaled air is lower than the temperature
of the body. As many animals, such as gazelles, inhale warm air, heat and
water vapour from the nasal passages evaporate, cooling the nose and
the blood within it. The cool venous blood passes close to and cools the
warm arterial blood traveling to the brain. If the brain does not require
cooling, the venous blood returns to the heart by another route. The
nasal passages also cool the warm, saturated air from the lungs so that
water condenses in the nose and is reabsorbed rather than lost to the
environment.

ph

The relative acidity or alkalinity of a solution is reported by the pH scale,

which is a measure of the concentration of hydrogen ions in solution.
Neutral solutions have a pH of 7. A pH of less than 7 denotes acidity (an
increased hydrogen ion concentration), and above 7 alkalinity (a
decreased hydrogen ion concentration). Many important molecular
processes within the cells of organisms occur within a very narrow range
of pH. Thus, maintenance of internal pH by homeostatic mechanisms is
vital for cells to function properly. Although pH may differ locally within
an organism, most tissues are within one pH unit of neutral. Because
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aquatic organisms generally have somewhat permeable skins or
respiratory exchange surfaces, external conditions can influence internal
pH. These organisms may accomplish the extremely important task of
regulating internal pH by exchanging hydrogen ions for other ions, such
as sodium or bicarbonate, with the environment.

The pH of naturally occurring waters can range from very acidic

conditions of about 3 in peat swamps to very alkaline conditions of about
9 in alkaline lakes. Naturally acidic water may result from the presence
of organic acids, as is the case in a peat swamp, or from geologic
conditions such as sulfur deposits associated with volcanic activity.
Naturally occurring alkaline waters usually result from inorganic sources.
Most organisms are unable to live in conditions of extreme alkalinity or
acidity.

Salinity

The term salinity refers to the amount of dissolved salts that are present
in water. Sodium and chloride are the predominant ions in seawater, and
the concentrations of magnesium, calcium, and sulfate ions are also
substantial. Naturally occurring waters vary in salinity from the almost
pure water, devoid of salts, in snowmelt to the saturated solutions in salt
lakes such as the Dead Sea. Salinity in the oceans is constant but is more
variable along the coast where seawater is diluted with freshwater
from runoff or from the emptying of rivers. This brackish water forms a
barrier separating marine and freshwater organisms.

The cells of organisms also contain solutions of dissolved ions, but the
range of salinity that occurs in tissues is more narrow than the range that
occurs in nature. Although a minimum number of ions must be present
in the cytoplasm for the cell to function properly, excessive
concentrations of ions will impair cellular functioning. Organisms that
live in aquatic environments and whose integument is permeable to
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water, therefore, must be able to contend with osmotic pressure. This
pressure arises if two solutions of unequal solute concentration exist on
either side of a semipermeable membrane such as the skin. Water from
the solution with a lower solute concentration will cross the membrane
diluting the more highly concentrated solution until both concentrations
are equalized. If the salt concentration of an animal’s body fluids is
higher than that of the surrounding environment, the osmotic pressure
will cause water to diffuse through the skin until the concentrations are
equal unless some mechanism prevents this from happening.

Many marine invertebrates have the same osmotic pressure as

seawater. When the salt concentration of their surroundings changes,
however, they must be able to adjust. Two means of contending with
this situation are employed, and, depending on how they regulate the
salt concentrations of their tissues, organisms are classified as
osmoregulators or osmoconformers. The osmotic concentration of the
body fluids of an osmoconformer changes to match that of its external
environment, whereas an osmoregulator controls the osmotic
concentration of its body fluids, keeping them constant in spite of
external alterations. Aquatic organisms that can tolerate a wide range of
external ion concentrations are called euryhaline; those that have a
limited tolerance are called stenohaline.

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Even if aquatic organisms have an integument that is relatively
impermeable to water, as well as to small inorganic ions, their
respiratory exchange surfaces are permeable. Hence, organisms
occurring in water that has a lower solute concentration than their
tissues (e.g., trout in mountain streams) will constantly lose ions to the
environment as water flows into their tissues. In contrast, organisms in
salty environments face a constant loss of water and an influx of ions.
Many mechanisms have evolved that deal with these problems. Because
water cannot be readily pumped across cell membranes, salinity balance
is usually maintained by actively transporting inorganic ions, usually
sodium and chloride. This process consumes energy and can usurp a
large portion of the energy budget of animals in very saline
environments. In marine fish, gill cells pump ions out of the body into the
sea, while in freshwater fish gill cells pump ions in the opposite direction.
Passive water loss in marine fish is compensated primarily in one of two
ways. Most bony fish drink copiously and excrete salt across the gills,
while the majority of sharks artificially elevate the salt concentration of
their tissues above that of seawater with urea and other organic

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molecules, allowing water to slowly and passively enter the body.
Through their food and across their gills, freshwater fish replenish most
of the ions they lose. They also produce large quantities of very dilute
urine to excrete excess water that diffuses into their bodies (Figure 8).

osmotic regulation
Figure 8: Osmotic regulation in freshwater and marine teleost fish.
Encyclopædia Britannica, Inc.

Water currents

The flow of water presents special problems for aquatic organisms. Flow
is associated with rivers, oceanic currents, and waves and can be laminar
(streamlined) or turbulent. Many organisms are specialized to live in
flowing environments; the main obstacle to this lifestyle is the constant
threat of being washed away. Both plants and animals have evolved
mechanisms that help to anchor them to the substratum in flowing water
(e.g., the holdfast of kelp or the byssus threads of mussels). If anchorage
can be assured, there are many advantages to living in this environment.
Flowing water generally is well oxygenated, and the supply is continuous;

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nutrients and food are constantly replenished as well. The very
precariousness of the environment also affords some protection from
predation because the number of predators that make this type
of habitat home is limited.

Pressure

Atmospheric pressure

Variations in atmospheric pressure can present special problems for

the respiratory systems of animals because atmospheric pressure affects
the exchange of oxygen and carbon dioxide that occurs during animal
respiration. Normal atmospheric pressure at sea level is the total
pressure that a column of air above the surface of Earth exerts (760
millimetres of mercury, or 1 atmosphere). The total pressure is the sum
of the pressures that each gas—mainly nitrogen, oxygen,
and carbon dioxide—would exert alone (the partial pressure of that
gas; see respiration: The gases in the environment). As an animal
breathes, oxygen moves from the environment across the respiratory
surfaces into the blood; carbon dioxide moves in the reverse direction.
This process occurs primarily by passive diffusion; each gas moves from
an area of greater to lesser partial pressure, driven by the differential
that exists across the respiratory surface. At higher altitudes, where the
atmospheric pressure is lower, the partial pressure of oxygen is also
lower. The partial pressure differential of oxygen, therefore, is also
lower, and the organism effectively receives less oxygen when it
breathes, even though the percentage of oxygen in the air remains
constant. This lack of oxygen is why humans carry oxygen when
ascending to high altitudes. Humans who live in mountainous regions,
however, can become acclimatized to the lowered availability of oxygen,
and certain animals such as llamas have adaptations of the blood that
allow them to live at high altitudes. Birds have very efficient lungs, and

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many apparently have no problems flying to high altitudes, even for
extended flights.

Hydrostatic pressure.

Because air and water have vastly different densities, the pressures
experienced in terrestrial and aquatic habitats differ markedly. A column
of water, so much denser than air, exerts a greater amount of pressure
than a column of air. With each 10-metre (32.8-foot) increase in depth,
there is an increase in hydrostatic pressure equivalent to
one atmosphere. Mean ocean depth is about 3,800 metres and has a
pressure of about 380 atmospheres. To surmount this environmental
challenge, animals that live at great depths lack air compartments such
as lungs or swim bladders. Surface-dwelling animals that dive to great
depths meet this challenge differently. As pressure increases during a
dive, air compartments compress, returning to their former volume
when the animal surfaces. Air is forced into the trachea, bronchi, and
bronchioles, where no gas uptake occurs. Thus, the increased pressure
cannot drive more gases into the bloodstream, and, as the animal rises,
it does not experience the “bends” (decompression sickness resulting
from a rapid reduction of air pressure). In contrast, sea snakes avoid the
bends by excreting nitrogen across the skin to offset the uptake of this
gas from the lungs.

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The 10 Percent Rule
https://youtu.be/ScizkxMlEOM

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