SEED COURSE

Unit 1- Limnology- An introduction

1.1.1 Introduction
Civilizations have depended on water bodies such as lakes, reservoirs, rivers and wetlands.
Water is essential not only to sustain human life but also to support the activities that form the
basis for thriving economics. Though the water resources are essential to human societies who
could pollute and degrade and limiting their beneficial uses. Agriculture, mining, urban
development and other activities can pose risks to freshwater bodies and hence steps have to
be taken to reduce these risk factors.
Risk analysis requires knowledge of how human land use affects physical, chemical and
biological characters of the aquatic systems. One of the critical areas required to understand
how human actions and natural processes affect lakes, reservoirs, rivers and wetlands is the
science called Limnology. It is a multidisciplinary science that integrates the basic sciences
(Biology, Chemistry, Physics and Geology) in order to study inland waters as complex
ecological systems.

Definition
The term Limnology is derived from Greek word; Limne means lake and logos means
knowledge. Limnology is often regarded as a division of ecology or environmental science. It is
however, defined as “the study of inland waters” (running and standing waters fresh and some
times saline; natural or man made). This includes the study of lakes, ponds, rivers, reservoirs,
swamps, streams, wet lands, bogs, marshes etc. Hence, it is commonly defined as that branch
of science which deals with biological productivity of inland waters and with all the causal
influences which determine it (Welch, 1963).
Biological productivity, as used in this definition, includes its qualitative and quantitative features
and its actual and potential aspects. Under the term inland waters are included all kinds or types
of water – running or standing; fresh, salt or other physicochemical composition which are
wholly or almost completely included within the land masses. Causal influences involve various
factors – physical, chemical, biological, meteorological etc which determine the character and
quantity of biological production.

History
The term Limnology was coined by Francois-Alphonse Forel (1841 – 1912) who established the
field with his studies on Lake Geneva. Interest in the discipline rapidly expanded and in 1922
August Thienemann (a German Zoologist) and Einar Naumann (a Swedish Botanist) co-
founded the International Society of Limnology (SIL, for originally Societas Internalis
Limnologiae). Forel’s original definition of limnology, oceanography of lakes was expanded to
encompass the study of all inland waters.
Welch (1935) conceived the problem of “Biological productivity” as the central theme of
Limnology. He defined Limnology as that branch of science which deals with all causal
influences which determine it. According to Schwoerbel (1987), Limnology is the science of
inland waters viewed as ecosystems together with their structures, materials and energy
balance. Kiihnelt (1960) considered limnology as a sub set of ecology along with
“Oceanography” (which is concerned with marine ecosystem) and “Epheirology” (which deals
with terrestrial habitats). In short, Limnology is the study of all aquatic systems including lakes,
wetlands, marshes, bogs, ponds, reservoirs, streams, rivers etc. with regard to their physical
chemical and biological characteristics.
In addition to the above, certain other terms, like Hydrobiology, Freshwater Biology, Aquatic
Biology, Aquatic Ecology etc, are sometimes loosely used as synonymous to the word
'Limnology'. But, most of these terms are names under which a diverse variety of subject matter
is included and only a part of it is limnological in nature.

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1.1.2 Development of Limnology

Initiation of works in the field of Limnology could be traced back to Aristotle (384-322 BC) which
consisted mostly of strange mixtures of facts and fiction with little scientific value followed by
observation of certain conspicuous freshwater phenomena. However, significant contributions of
strictly limnological nature began to appear about a thousand nine hundred years after Aristotle
with the description related to habitat, habit and life history of certain fishes, insects and aquatic
macrophytes. But, most of these were isolated accumulation of correlated observations, few of
which could be used by modern limnologists mostly for historical purposes.
Since the initiation of optics with Euclid (2000 BC) and later with the invention of microscope,
there has been significant development in the field of aquatic biology and limnology because it
has not only opened the door to the whole world of microscopic organisms, but also provided
with a new and effective means of studying the various higher types of life in water. This was
followed by description of minute aquatic organisms by Anton Van Leewenhook (1632 – 1723),
the pioneering classification of microscopic organisms by the Danish biologist, Otto Friedrich
Muller (1786), publication of the Treatise, “Infus Animalcules” by Ehrenberg (1838) which marks
the beginning of those advances in knowledge which occurred in the 20th century.
Peter Erasmus Muller is credited with laying the foundation stone of limnological study. Anton
Fritsch could be considered as the pioneer in lacustrine limnology for his work on lakes in the
Bohemian Forest and F. Simony (1850) is regarded, sometimes as the founder of Limnology for
his discovery of thermal stratification. However, it remained practically everything for F.A. Forel
(1841- 1912), a professor in the University of Lausanne, Switzerland, to recognize the real
biological opportunity of lake investigations and the science of limnology is indebted to him for
his comprehensive vision and complete anticipation about the future of this subject. He is
regarded as the Founder and father of Modern Limnology for his 110 publications (Chumley,
1910). It was he who took the decisive step forward from hydrobiology to limnology through his
investigations in Lake Geneva, not only from the biological point of view but also from physical
and chemical stand points, thereby formulating the concept of lake types. In addition the design
of his first programme for limnological investigations in freshwater and its subsequent execution
turned out to be a model for future researches.
Forel’s work paved the way for establishment of Limnological Society in 1887 as a component
of Swiss Natural history Society (in order to promote limnological works) and later the
International Commission of limnology was established in 1890). In brief, the History of
limnology could be dated back to approximately 100 years. Although certain preliminary studies
has been done on the habits, nutrition, movement, behavior etc., on certain aquatic organisms
by different workers during the 17th and 18th centuries, these were mostly hydrobiological
works and not limnological. True studies on the relationship of biota to freshwater could be
treated as initiated from Junge (1885) and Forbes (1897) who were the first to treat the native
waters as microcosm.
Gaarder and Gran (1927) made pioneering attempts at measuring the photoautotrophic
production (primary production) by quantitative determination of oxygen produced during
photosynthesis. Later the direct measurement of carbon assimilation in the water bodies was
achieved in 1952 using radio-carbon method (Steemann and Nielsen, 1952). The estimation of
trophic dynamics concept having regard to the biomass, material turn over and energy transport
along the food chain by Lindeman (1942) not only revolutionized the field of general ecology,

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but also gave a new direction to Limnology (Cook,1977).

Early freshwater investigations

In 1870, Simson, published a short account of the deep water fauna of Lake Michigan. Smith
and Verrill (1871) made deep water dredging in Lake Superior and published on the
invertebrates collected. In 1886, the Allis Lake laboratory, a privately supported institution and
said to be the first freshwater biological station in America, was established at Milwaukee,
Wisconsin, but its life was brief and none of its work was concerned with the general biology of
the Great lakes. In the meantime, interested workers were giving attention to some of the
smaller inland lakes. Forbes made a study of certain high lakes of the Rocky Mountains and
published only on biological information concerning lakes in western United States. During the
decade of 1890 -1900, important freshwater biological stations have been found viz, (1) the
University of Minnesota at Gull Lake, Minnesota, 1893; (2) the University of Illinois on the Illinois
River, 1894; (3) the University of Indiana at Turkey Lake, Indiana, 1895.
The stimuli of scientific interest and of the necessities of public health brought about the
initiation of systematic surveys of water supplies and of water systems in general, the
Massachusetts State Board of health taking the lead in about 1887. Subsequently, similar work
was undertaken by various municipal and government departments, all of which contributed,
directly and indirectly to the sum total of limnological information. Phenomenal progress of the
general subject of ecology inevitably had a constructive influence on limnology and because of
its many ramifications limnology has likewise profited from simultaneous advances of other
sciences.

1.2.1 Inland waters
The inland waters which include both fresh water masses and estuarine waters of varying salt
content are clearly distinguishable from the salt waters of the oceans. The inland water masses
are discrete and being isolated within the specific land area, acquire the characteristic chemical
composition of the land, by exchange between soil and water. The oceanic water on the other
hand is open and mixing together by wind action and currents and therefore more
homogeneous in chemical composition. However, the land water exchange is limited to coastal
areas. The estuarine waters are mixtures of sea and freshwater, but with the higher content of
salts in the sea water (150 – 200 times that of freshwater), are dominated by the sea water
effects.
According to Hutchinson (1959), limnology is the large variety, individual and groups of inland
water bodies, the diversity being caused by the diversity of their origin as well as by the diversity
of their chemistry and biology.

Types of inland water

Frey (1960) has classified inland waters in three different ways viz, depending on whether the
water is stationery or flowing, depending on whether the water mass is natural or artificial and
permanent / temporary.
I a. Flowing waters (Lotic waters)
These include creeks, streams and rivers mentioned in that sequence because of their
sequence of succession also in the same order, through the natural processes of lengthening
and widening of running waters. In these, there is continuous current of water in one direction.

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The organisms inhabiting these waters have complexity of adaptation towards the increase in
water current speed. It includes all forms of inland waters in which the entire body of water
moves continuously in a definite direction. The sequence of genesis is brooks, rivulets, channels
and rivers. 
b. Standing waters (Lentic waters)
Here, water current is not a major ecological factor; unlike in the lotic series lakes, ponds and
swamps form the lentic series. The sequence indicates the natural evolution of water masses as
well lake may either be productive or non-productive, when they are referred to as eutrophic or
oligotrophic respectively. It includes all forms of inland waters – lakes, ponds, swamps and their
various integrades in which the water does not flow continuously in definite directions.
Essentially, the water remains standing, though a certain amount of water movement may
occur, such as wave action, internal currents or flow of water in the vicinity of inlets and outlets. 
The sequence of genesis is as follows Lake – pond – swamp.
a. Natural bodies of water
Certain parts of the world are endowed with an abundance of natural waters serving human
needs.
b. Artificial bodies of water
According to man’s needs water bodies are created artificially. It includes ponds, wells, tanks
reservoirs etc.

i. Ponds 
In India, even from ancient times, large ponds and wells exist serving for drinking water and also
for irrigation purposes.
Types of ponds 
Based on seasonal duration ponds can be classified into two types. 
1. Temporary ponds 
2. Seasonal ponds 
Temporary ponds divided into three types, 
1. Vernal ponds: Water exists only in spring season. 
2. Vernal Autumnal pond: Water exists in those ponds during spring and autumn and they dry in
summer. 
3. Aestival ponds: Water persists in these ponds throughout the season but it freezes during
winter. 
Permanent ponds
Water persist in these ponds throughout the season but it freezer in winter. 
i. Reservoirs
Rivers are blocked and reservoirs or artificial lakes are developed. These serve in generating
hydroelectric power, irrigation, fish production and recreation. These also help in flood control. 
ii. Tanks 
In India there are both perennial and temporary tanks. There are some in which water remains
for 6-9 months duration, called long seasonal tanks and in some, water remains for less than 6
months, referred to as short seasonal tanks.

III a. Permanent waters

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In most parts of the world where there is precipitation exceeds, the evaporation and seepage
loss the waters in rivers, lakes and ponds and are termed as permanent waters.
b. Temporary waters
Evaporation loss of water is more than the precipitation gains, as in all arid areas of the world,
the water bodies dry up usually during summer. In high latitude (30-50o), when the area is
desert the rivers and streams drain into permanent or temporary lakes. These lakes have salt
differing from that of sea, the salt washed down to the lakes being predominantly potassium and
sodium carbonates and magnesium and sodium sulphates and not sodium chloride. By
evaporation at times salt concentrations in these waters exceeds that of the sea. (eg. Great salt
lakes and Dead sea). The salt concentration of Dead sea is so high that there is no life in it.

1.2.2. Distribution of inland waters

Inland waters cover less than 2% of the earth’s surface, approximately 2.5 x 106 km2. About 20
lakes are extremely deep (in excess of 400 m). A significant portion of the world’s freshwater is
contained in lake. Some regions are very generously supplied with lakes and streams
particularly those regions once subjected to ancient glaciation. Canada and northern United
States possess an immense supply of lakes, among them the Great lakes, which constitute the
greatest body of freshwater on the globe. Portions of Europe are also noted for their generous
supply of lakes and streams. In certain regions, disappearance of inland waters during the dry
season forms the basis for special biological phenomena resulting from the intermittent
character of the environments.
In India, most of the wetlands including flood plain wetlands are situated in the eastern parts of
the country whereas, reservoirs and tanks have been created mainly for irrigation are distributed
throughout the country. Large rivers and streams are well distributed in the northern and eastern
region of the country. However, southern regions of the country also have a good number of
inland waters. Many of them are seasonal in nature. 

Dynamics of Lotic and Lentic environment 

In the lotic series, the tiny rivulet gradually deepens, widens its bed, and cuts back at its head,
thus in time extending its length and increasing its cross section to that size which justifies the
designation of brook. This process continues by the same general type of action, ultimately
producing a creek and then finally a river, with all of the integrating conditions produced in such
a gradual transformation.
Faunas occupying each of the different environments must accompany these migrations or
become adapted to the gradually altering conditions or they will become extinct. These
environmental migrations are very slow, in point of time, and give ample opportunity for the
characteristic organisms of particular environments to make the necessary responses. The
ultimate fate of any lotic series is the degradation of the land is the reduction of its bed to base
level.
In the lentic series, natural processes work toward extinction, mainly by the gradual filling of
basins.
Lake –> Pond –> Swamp
In larger lakes, natural filling takes much longer time, even many centuries also hence the filling
is primarily due to :
• Wind blown materials such as dust, sand and debris of various sorts. 

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• Sediments brought into a lake by inflowing streams and by incoming run-off water as it flows
down adjacent land slopes. 
• Wave action, cutting away exposed shores and depositing eroded material in lake basin. 
• Plants, particularly the higher aquatic plants which grow in shallow water, produce deposits of
organic matter. 
• Accumulating remains of animal life especially shells. 
Not all lakes become extinct by filling alone. Other process also contributes to this for example
an outlet may cut down its level at the point of exit from a basin, thus gradually draining the lake.
These stages in the extinction of standing waters result in a more or less definite, predictable
‘evolution of environment’ in the long run has a profound influence on the history and fate of
lake organisms.

Running waters (Lotic series)

There are many different kinds of running waters, several of them occurring, inter-connected,
within a single drainage system. The range covered within the series includes small trickles and
seepages, ditches, larger fast flowing streams and rivers, large slow flowing rivers and canals.
The flow characteristic of any running water system are also closely connected with the
geology, notably in the control exerted by the nature and structure of rock and soil formations,
and also in the relationship between the amount of ground water and surface water flowing
through the system. The basic flow pattern depends largely on the nature of this relationship.
Most of the water on the earth is in constant circulation within what is known as ‘hydrologic
cycle’. The energy utilized within the cycle comes mainly from the sun. Water evaporates from
both land and sea to be re-precipitated, usually somewhere else. On most part of the land,
precipitation exceeds evaporation, and run off towards the sea occurs.

Difference between running water and standing water

• Current : Unidirectional main current is found in running water but not in standing water.
• Depth: It is small in running water, more in standing water. 
• Condition of gradient from source to mouth : In running water, physical, chemical conditions
usually change from the source to the mouth and the difference in many factors may be great
between those extremes, but it is more homogeneous in standing water. 
• Water of the basin: Running water systems are very shallow and have long, complex narrow
channels, but standing water reach great depth, have broad basins. 
• Permanent removal of eroded and transported materials: Constant erosion is common in
running water and materials so removed are transported to distance. Erosion occurs in standing
water, but it is rarely severe, eroded materials and not carried far away but remains within the
same basins. 
• Absence of prolonged stagnation: Consequence of erosion and deposition, most of the running
waters tend to increase the length of their channels with age. In standing water materials
constantly being deposited tend to fill in the basin. 
• Physical factor: It is more important in running water and standing water. 
• Basic food materials: Most running water manufacture themselves little basic food, but depend
on the contribution from the surrounding land than the standing water. 

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1.3.1. Ponds, lakes, streams, river

Ponds:
Ponds are defined as small, shallow, inland standing water bodies, where rooted plants can
grow over most of the bottom. Ponds are mainly of three general classes, they are : 
i. Those which represent the pond stage in the extinction of previously existing lakes
ii. Those whose basins have never been large or deep (not preceded by a lake) but or for some
special reason, have persisted in the pond stage and
iii. Those whose basins are the results of man’s activities (excavations, quarries,
impoundments, etc.) 
Natural process alone are constantly forming new pond basins (cut-offs from streams solution
basins, beach ponds, and many others), some of which are never more than temporary ponds
from the beginning; others qualifying as permanent ponds at least for a period in their existence.

Classification of ponds 
With respect to seasonal duration, ponds are divided into two general classes
a. Permanent – those which contain some water the year round and
b. Temporary – those in which the basin contains water at certain times or seasons and
becomes dry at others.
Those which occur for a limited period in spring are called Vernal ponds
Those which contain water in spring, dry up during summer, and again contain water in the
autumn are called Vernal autumnal pondsand
Those which contain some water throughout the open season but freeze to the bottom in winter
have been called Aestival ponds.

Other classifications of ponds are as follows

Natural ponds
These are perennial shallow water bodies. When a stream shifts its position it leaves behind an
isolated body of standing water which forms the "Ox-Bow" pond. In limestone regions where
depressions are formed due to the solution of the underlying strata, the water gets accumulated
either by flood water or rainfall and natural ponds are formed. Sometimes the last remnant of a
lake whose basin has become filled progressively by sedimentation in course of time is
transformed into a pond. 
Artificial pond
Most of the fish ponds are semi artificial ponds. Some are constructed by erecting dams across
a stream or basin and their water level can be regulated by inflow and drainage where
pisciculture is practiced. Fish pond is a shallow body of water that can be drained completely. It
is often supplied by running water, but also by spring, ground or rain water.
Pools or Temporary Ponds
They occur in depressions in the ground either at the margin of glaciers where they fill with melt
water or in the vicinity of river bed, after the floods have receded. The water thus collected
usually is very shallow and measures maximum to a few feet only. Also prolonged rainfall may
form temporary small pools. All these pools dry up in some part of the year, and as such
organisms in these habitats must be able to survive in a dormant stage during dry periods and
be able to move in and out of the pools.

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General Characteristics of ponds

• Ponds are small, shallow standing water bodies.
• They have calm water 
• Have more vegetation
• Growth of plants can also found at the bottom
• They have outlet streams
• The movement of water is minimum
• They have slight wave action
• The average depth of water is 8 – 10 feet
• The temperature of the pond more or less changes with that of atmosphere
• Light penetrates up to the bottom

1.3.2. Lakes
Forel (1982) defined lake as a body of standing water and occupying basin and lacking
continuity with the sea. He also defined pond as a lake of small depth, and a swamp has been
defined as a pond of small depth and occupied by rooted vegetation. Carpenter (1928)
formulated that the true difference between lake and pond is depth and not area accordingly a
pond is a quiet body of water where floating vegetation extends to the middle of basin in which
the biota is very similar to littoral zone of lake.

Lake Morphology
The shape of a lake basin is largely determined by its mode of origin. The depth and contour of
lake bottom can be determined by lowering a weighted line or much more quickly with an echo
sounder. Physical structural components of lakes include their shape, distribution of light,
distribution of heat, and movement of water. The hydraulic retention time (time required for all
the water in the lake to pass through its outflow) is an important measure for lake pollution
studies and calculations of nutrient dynamics. The hydraulic retention time is mainly determined
by the interplay between inflow of water into the lake and the basin shape.

Lake Zonation
The following depth zones are recognized in lakes:
a. littoral zone extends from the shore just above the influence of waves and spray to a depth
where light is barely sufficient for rooted plants to grow.
b. photic (euphotic) zone is the lighted and usually well-mixed portion that extends from the
lake surface down to where the light level is 1% of that at the surface.
c.aphotic zone is positioned below the littoral and photic zones to bottom of the lake, where
light levels are too low for photosynthesis. Respiration occurs at all depths so the aphotic zone
is a region of oxygen consumption. This deep, unlit region is also known as the profundal zone.
d.compensation depth is the depth at which rates of photosynthesis and respiration are equal.
e.sublittoral zone, which is the deepest area of plant growth, is a transition between the littoral
and profundal zones.
f.pelagic zone (limnetic zone) is the surface water layer in offshore areas beyond the influence
of the shoreline.
Boundaries between these zones vary daily and seasonally with changing solar intensity and

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transparency of the water. There is a decrease in water transparency with algal blooms,
sediment inflows from rivers or shore erosion, and surface waves.

1.3.3. Streams and Rivers

A. STREAMS
Introduction 
Streams are zones where a rapid flow of shallow water produces a shearing stress on the
stream bed, resulting in a rocky or gravel substratum covered by fully oxygenated water.
Streams may vary in size from tiny rivulet to rivers. As time goes the stream may develop into
river or increase its size, whereas the size of reservoirs decreases as time passes. They are
more numerous in regions of abundant rain fall. They are temporary or permanent. Streams are
closely linked to their watersheds. The productivity of streams is often dependent on terrestrial
bases, grasses and other debris. The allocthonous materials contribute most of the food and
energy to the organisms in the stream. Benthic invertebrates like insect larvae constitute the
invertebrate fauna. True plankton are almost absent in streams, and are common only in deep
slow moving stretches of rivers. All biota in streams are influenced by the unidirectional current. 

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Physical conditions 
The annual change in stream temperature is 10 to 20°C. Although large rivers do not change in
temperature very much on a daily basis, a small unshaded stream may heat up to 10°C in a few
hours on a hot summer’s day and cool by the same amount at night. The temperature of most
streams is lowest in the upland and becomes gradually warmer in the lower reaches. 
The velocity of stream water varies with the landforms. In plains, streams are slow and sluggish
throughout their length. In mountain stretches the speed of water may be rapid.
Stream water has uniform temperature and the difference between the surface and bottom is
virtually negligible. The stream follows air temperatures more closely than lake waters and the
factors responsible are depth of water, current velocity, bottom material, temperature of entering
water, exposure to direct sunlight and degree of shading etc.
Extreme of turbidity occur in running water series and streams with rock beds the turbidity is
minimal.
Stream systems increase their length, width and depth with increasing age. This is in distinct
contrast to the reduction processes characteristic of all standing water units.
At any position along the course of a running water system, materials eroded at that point and
all materials suspended or dissolved at the level are transported downstream with no
opportunity to return. Interchanges of materials are more and have less depth than lakes.
Chemical conditions
The dissolved oxygen supply in uncontaminated stream is high at all levels often near
saturation. The polluted streams show low dissolved oxygen due to accumulation of organic
wastes. Stream which support more plants show diurnal variation of dissolved oxygen. The level
of dissolved oxygen is controlled by the slope of channel and mode of flow. 
Current in streams tends to keep the pH in uniform over considerable distances. It keeps any
acidity due to accumulating free CO2reduced. Streams waters do not develop the more intense
acidities.
The dissolved solids of streams are affected by their irregular discharges. Most streams and
rivers have maximum discharge during winter rains, particulate matters, nutrients like
phosphate, iron and nitrate are transported to different parts by the flow of the streams. Streams
fed by springs have more constant nutrients.

B. RIVER

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River is said to be a natural stream of water usually fresh water flowing towards an ocean. In
some cases river flows into the ground or dries up completely before reaching another body of
water. Usually larger streams are called rivers while smaller streams are called creeks, brooks,
rivulets, rills, and many other terms. 
A river is a component of the hydrological cycle. The water within a river is generally collected
from precipitation, through surface run off, ground water recharge and release of stored water in
natural reservoirs such as glacier.
Topography
The water in a river is usually confined to a channel, made up of stream bed between banks. In
larger rivers there is also a wider floodplain shaped by flood waters over-topping the channel.
Flood plains may be very wide in relation to the length of river channel. This distinction between
river channel and floodplain can be indistinct especially in urban areas where the floodplain of a
river channel can become greatly developed by housing and industry. 
Ecology
The flora and fauna of rivers use the aquatic habitats available, from torrential waterfalls through
to lowland mires although many organisms are restricted to the freshwaters in rivers eg salmon
and Hilsa.
Flooding
Flooding is a natural part of river cycle. The majority of the erosion of the river channels and the
erosion and deposition on the associated flood plain occur during flood stage. Human activity,
however has upset the natural way flooding occurs by walling of rivers set straight their courses
and by draining of natural wetland.

Unit 2- Lakes
2.1.1 Lakes
Lakes origin
Lake is defined as a large body of standing water occupying a basin which does not have any
connection with sea. Approximately 1% of water is found in lakes, but the renewal time is much
more rapid than the ocean.
Classification of lakes on the basis of their origin
1.Tectonic lakes
These are formed in basins created by movements of the earth’s crust by different processes.

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2.Crater lakes (Volcanic lakes)

In many cases when a volcano becomes extinct, its hollow interior is filled with water by
precipitation or by percolation. They are near circular or sometimes elliptical in outline. eg,
Crater lake in Oregon (USA).

3.Glacial lakes
Most common lakes are originated due to erosion and deposition associated with glacial ice
movements eg. Finger lakes of New York.

4.Basal rock dissolution lake

These are formed by the slow dissolving of soluble rock (Calcium carbonate) by water eg, clear
lake in California and Deep lake Florida (USA).

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5.Oxbow lakes
An oxbow is a crescent-shaped lake lying alongside of meandering streams or rivers in the floor
of a valley. The oxbow lake is created over time as erosion and deposits of soil change the
river's course. On the inside of the loop, the river travels more slowly leading to deposition of
silt. Water on the outside edges tends to flow faster, which erodes the banks making the
meander even wider. When the streams bend and are cut off from the main stream flow, an
oxbow lake results. Such lakes may be entirely cut off and become totally lentic or a little flow
may persist seasonally at floods. eg, Dal and Woolar lakes of Kashmir.

6.Fluvial lakes
These lakes are formed by river activity.

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7.Aeolian lakes
These are formed by the wind activity in arid regions which may erode with broken rocks or
redistribute sand which are generally temporary.

8.Shoreline lakes
Created by irregularities or inundation along the coastline of large lakes which usually a result of
long shore currents.

9.Reservoir lakes
These are man made lakes formed by the construction of dams across the streams eg,
Thungabhadra reservoir in South India, Bhakra-Nangal reservoir in north western part of India
(Anthropogenic lakes).

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10.Bog lakes
Bogs are best developed in the north temperate glaciated region when precipitation is abundant
throughout the year, atmospheric humidity is great. Soil temperature is low, evaporation is
reduced, and run-off water is minimum having abundant growth of plants. A typical bog lake is
defined as an area of open water surrounded either wholly or by part of true margins.
Possessing peat deposits about the margins or in the bottom usually with a false bottom of very
finely divided flocculent vegetable matter.

11.Salt lakes
There are many salt lakes throughout the world, much as saline lake waters as freshwater
lakes. When climate change become drier or geological events change drainage basins, the
annual flows into lake may be greatly reduced. The lake may seem to have a significant outflow
and become a terminal or sink lake. The salts from the flowing stream are concentrated by
evaporation and are no longer flushed out through outflow eventually the lake may dry
completely. In dry climates the lakes gradually become salty.
Lake Bonneville and Lahontau of south western United States, the Great Salt Lake, Utah and
lake Walker and Pyramid in Wiveda are other examples.
Salt lakes also found in areas of drier climates such as Australia, South America, East Africa,
Antarctica, Russia and dry northern side of Himalayas, South and North East America.

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2.1.2 Thermal Classification of lakes 

According to Hutchinson (1957), following are the classification of lakes based on changes in
temperature of surface water.
a. Amictic: No mixing of bottom and top water; lakes insulated or protected by ice-corer, there
is no effect of weather or external factors.
b. Monomictic: One mixing of the two waters during the year (most deep lakes of the world).
c. Cold monomictic: Water here at any depth never exceeds 4°C; they are ice-bound or ice-
covered only in winter; there are inverse thermal stratification top waters 0°C and bottom waters
4°C (since water at 4°C is heaviest); only one mixing at temperatures not more than 4°C in
spring / summer eg, Polar lakes.
d. Warm monomictic: Temperature of water never falls below 4°C at any depth. Direct thermal
stratification top waters 10 - 20°C and bottom waters 8 - 4°C; only one mixing in a year in a
winter eg, Most subtropical deep lakes.
e. Ploymictic : Mixing is continuous, but occurs only at low temperatures.

Size of lake
Lakes differ in area from those ranges from a pond to those of great size. Lake Superior, the
largest body of freshwater flow has an area of more than 49,600 km2. The Caspian sea with an
area of 2,72,000 km2 is sometimes considered as having the quality of lake. 
Lake Chad in Africa has 64,000 km2 during wet season, but is reduced to 9,600 km2 in the dry
season. Ten of the large lakes in America including Great lakes have an combined area of
about 2,03,200 km2. However the number of lakes whose area exceeds more than 8,000 km2 is
insignificant when compared to many thousands of lakes of lesser magnitude of 11,000 or more
lakes or ponds in Michigan.
Depth of lake 
Lakes vary in depth but even the deepest lake will never approaches the depth of ocean. It is
important to note that the lake Baikal has a greatest depth contains about 20% of the total
volume of freshwater and it is also the deepest known lake with a maximum depth of 1620 m. In
North America, Crater lake in Oregon is about 608m. Lake Tahoe is 487m, Lake Chelan
(Washington) 457m. Seneka lake 188m, Lake Superior 393m, Lake Michigan 281m, Lake
Huron 228m, Lake Ontario 273m, Lake Erie 64m, and the last 5 lakes constitute Great lakes of
America.
Lake margin
Nature of margin

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The line of demarcation between land and water is the margin of the lake which depends upon a
number of circumstances. 
Shore dynamics
Water is in some form of motion ranging from gentle to violent. It has a great potentiality to
cause changes on the shore against which it beats. In lake, the wave action is the principle form
of water movement that cause shore changes. Modification of the original shore line has been
accomplished by two main phenomenon such as shore cutting and shore building.
Lake bottom
The term lake bottom includes all part of bottom of lake basin from the water edge to the
deepest region. All lakes of particular region may have the same origin, may have begun the
history with same material and exist at the same climatic condition, yet the bottom material may
often be different in different lakes. The kind of bottom deposits and the rate of deposition may
depend upon the local circumstances. The nature of bottom deposits determines the biological
productivity. The principle sources of bottom materials are 
i. Bodies of plankton organisms which die and sink.
ii. Plant and animal remains.
iii. Organic and inorganic materials.
iv. Silt, clay and similar materials.
v. Marl or CaCO3 precipitated produced by plants and animals.
vi. Remains of floating blanket algae.
Diversity of lake
Though all the lakes appear to be similar, there may be differences in colour, taste, hardness,
turbidity and aquatic animals and plants. With the knowledge and the modern methods of
environmental analyses, the lakes posses physical, chemical and biological diversity. According
to a great diversity, lake may be stated into different forms as under : 
a. Large, medium or small.
b. Deep or shallow.
c. Protected or unprotected.
d. With or without tributaries and outlets.
e. Fresh, brackish or salt.
f. Turbid or clear.
g. Acid, natural or alkaline
h. Hard, medium or soft.
i. Surrounded by bog, swamp, forest or open shores.
j. High or low in dissolved content.
k. With or without stagnation zones.
l. With mud, muck or mucky sand or false bottom.
m. With high, medium / low biological productivity.
n. With / without vegetation beds.
o. Young, mature and senescent.

2.2.1. Famous lakes

Indian lakes
Almost every region of the country is dwelt by several of lakes that add great charm to their
natural characteristics. Rajasthan and Himachal Pradesh, North-Western state and Northern

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state of India, respectively are undoubtedly in possession of larger number of lakes than
anywhere else in the country. 
Dal lake-Jammu-Kashmir
Dal Lake is a lake in Srinagar in Jammu and Kashmir. The urban lake, which is the second
largest in the state, is integral to tourism and recreation in Kashmir and is nicknamed the "Jewel
in the crown of Kashmir" or "Srinagar's Jewel". The lake is also an important source for
commercial operations in fishing and water plant harvesting.
Location : Srinagar, Jammu and Kashmir
Lake type : Warm monomictic
Surface area: 18–22 square km
Average depth : 1.42 m (4.7 ft)
Catchment area : 316 square km (122 sq mi)
Max. length : 7.44 km (4.62 mi)
Max. width : 3.5 km (2.2 mi)

Hebbal Lake- Karnataka

Hebbal Lake is located in the north of Bangalore at the mouth of National Highway 7, along the
junction of Bellary road and the outer ring road. It was one of the three lakes created in 1537 by
Kempe Gowda. Like most lakes or "tanks" in the Bangalore region it was formed by the
damming natural valley systems by the construction of bunds. The spread of the lake in a study
in 2000 was found to be 75 ha with plans for extending it to make up 143 ha. The catchment
area of the lake was found to be 3750 ha. In 1974 the lake area was 77.95 ha and in 1998 it
was 57.75 ha. Based on the rainfall of the region, the annual catchment was estimated at 15.2
million cubic metres with 3.04 million cubic metres during the Northeast Monsoon, 10.12 million

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cubic metres during the Southwest Monsoon and 3.28 million cubic metres in the dry season.
The storage capacity of the lake was estimated in 2000 to be 2.38 million cubic metres with
desilting raising it to 4.07 million cubic metres. Sewage inflow into the lake has altered the
chemistry and biology of the lake.

Powai Lake-Maharasthra
Powai Lake is an artificial lake, situated in the northern suburb of Mumbai, in the Powai valley
located downstream of the Vihar Lake on the Mithi River. The city suburb called Powai, shares
its name with the lake. Population around the lake has substantially increased over the years.
When it was built, the lake had a water spread area of about 2.1 square kilometres (520 acres)
and the depth varied from about 3 metres (9.8 ft) (at the periphery) to 12 metres (39 ft) at its
deepest. The Powai Lake has gone through many stages of water quality degradation. The lake
water which used to supply to Mumbai for drinking water has been declared unfit to drink. The
Lake still remains a tourist attraction.
Location : Mumbai 
Catchment area : 6.61 km2 
Max. depth :12 m
Surface elevation : 58.5 m (191.93 ft)
Settlements : Powai 

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Loktak lake-Manipur
Loktak Lake, the largest freshwater lake in northeastern India, also called the only Floating lake
in the world due to the floating phumdis (heterogeneous mass of vegetation, soil, and organic
matters at various stages of decomposition) on it, is located near Moirang in Manipur state,
India. The etymology of Loktak is lok = "stream" and tak = "the end". This ancient lake plays an
important role in the economy of Manipur. It serves as a source of water for hydropower
generation, irrigation and drinking water supply. The lake is also a source of livelihood for the
rural fisherman who lives in the surrounding areas and on phumdis, also known as
“phumshongs. Considering the ecological status and its biodiversity values, the lake was initially
designated as a wetland of international importance under the Ramsar Convention on March
23, 1990.
Location : Manipur
Lake type : Fresh water (lentic)
Primary inflows : Manipur river and many small rivulets
Primary outflows : Through barrage for hydropower generation, irrigation, and water supply
Catchment area : 980 km2 (380 sq m)

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Chilka Lake - Orrisa

Chilka (Chilika) lake is a brackish water lagoon, spread over the Puri, Khurda and Ganjam
districts of Orissa state on the east coast of India, at the mouth of the Daya River, flowing into
the Bay of Bengal. It is the largest coastal lagoon in India and the second largest lagoon in the
World. It is the largest wintering ground for migratory birds on the Indian sub-continent. The lake
is home to a number of threatened species of plants and animals. The lake is an ecosystem
with large fishery resources. It sustains more than 150,000 fisher–folk living in 132 villages on
the shore and islands. Microalgae, marine seaweeds, sea grasses, fishes and crabs also
flourish in the brackish water of the Chilika Lagoon.
Lake type : Brackish
Primary inflows : 35 streams including the Bhargavi, Daya, Makra, Malaguni and Nuna rivers
Primary outflows : Bay of Bengal
Catchment area : 3,560 km2

Hussian Sagar - Andhra Pradesh

Hussain Sagar (Hyderabad, India) was built by Hazrat Hussain Shah Wali in 1562, during the
rule of Ibrahim Quli Qutb Shah. It was a lake of 24 square kilo metres built on a tributary of the
River Musi to meet the water and irrigation needs of the city. There is a large monolithic statue
of the Gautam Buddha in the middle of the lake which was erected in 1992.
Location: Hyderabad 
Lake type: artificial lake 
Max. depth: 32 ft
Surface elevation: 1,759 ft

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Brahma Sarovar - Haryana

Brahma Sarovar is a water tank sacred to the Dharmic religions in Thanesar, in the state of
Haryana in North India. Dharmic religions lay emphasis on taking bath for internal and external
purity. 
Max. width : 1,800 ft (550 m)
Surface area : 1,400 ft (430 m)

Vembanad Lake - Kerala

Vembanad Lake (Vembanad Kayal or Vembanad Kol) is India's longest lake, and is the largest

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lake in the state of Kerala. It is also one of the largest lakes in India.
Max. length : 96.5 km
Max. width : 14 km
Surface area : 1512 km2
Max. depth :12 m

Upper Lake - Madhya Pradesh

Upper Lake, is the largest artificial lake in Asia which lies on the Western side of the capital city
of Madhya Pradesh, Bhopal. It is a major source of drinkable water for the residents of the city,
serving around 40% of the residents with nearly 30 million gallons per day.
Location : Madhya Pradesh, Bhopal 
Primary inflows : Kolans River
Catchment area : 361 km²
Surface area : 31 km² 

Kodaikanal Lake - Tamil Nadu

Kodaikanal Lake, also known as Kodai Lake is a manmade lake located in the Kodaikanal city in
Dindigul district in Tamil Nadu, India. The lake is said to be Kodaikanal's most popular
geographic landmark and tourist attraction. Over the years a boat club, boathouse and boat
service for the public and tourists has become fully functional and is of aesthetic significance for
tourism. Boat Pageant and Flower Shows is a regular feature in the summer season which
attracts tourists.

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Location : Kodaikanal, Dindigul district, Tamil Nadu 

Lake type : Fresh water
Surface area : 24 ha (60 acres)
Average depth : 3 m (9.7 ft)

Pushkar Lake- Rajasthan

Pushkar is an artificial lake located in the state of Rajasthan in India. It is situated near the
Pushkar town in the district of Ajmer. The lake was constructed in the 12th century with the
establishment of the dam across the headwaters of the Luni river. The pious Pushkar Lake is
regarded as the sacred lake among the Hindus in India.

Osman Sagar Lake

Popularly known as the 'Gandipet', Osman Sagar Lake is the man made lake created by the
dam across the Isa, a tributary of the river Musi. It is the main source of water supply to the twin
cities of Hyderabad and Secunderabad.

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Bhimtal Lake 
Located 22 km from Nainital, and this lake is named after the second Pandava called Bhima of
the famous epic Mahabharata. It is one of the largest lakes in the Nainital and the second
largest lake in Kumaoun. The lake provides the excellent opportunity for boating, fishing and
angling.

Roopkund Lake
Roopkund Lake lies in the Chamoli district of Uttranchal at the height of 5029 meter. The lake
provides the stunning view of the Trishul peak (7122 meter) and due to its less depth it also
known as the shallow lake. 

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2.3.1. Lakes of the world

Largest by continent
The largest lakes (surface area) by continent are:
• Australia - Lake Eyre (salt lake)
• Africa - Lake Victoria, also the third-largest freshwater lake on Earth. It is one of the Great
Lakes of Africa.
• Antarctica - Lake Vostok (sub-glacial)
• Asia - Lake Baikal (if the Caspian Sea is considered a lake, it is the largest in Eurasia, but is
divided between the two geographic continents)
• Oceania - Lake Eyre when filled; the largest permanent (and freshwater) lake in Oceania is
Lake Taupo.
• Europe - Lake Ladoga, followed by Lake Onega, both located in northwestern Russia.
• North America - Lake Michigan-Huron, which is hydrologically a single lake. However, lakes
Huron and Michigan are often considered separate lakes, in which case Lake Superior would be
the largest.
• South America - Lake Titicaca, which is also the highest navigable body of water on Earth at
3,821 m above sea level. The much larger Lake Maracaibo is considered by some to be the
second-oldest lake on Earth, but since it lies at sea level and nowadays is a contiguous body of
water with the sea, others consider that it has turned into a bay.

Notable lakes
• Lake Michigan-Huron is the largest lake by surface area 117,350 km². It also has the longest
lake coastline in the world: 8,790 km. Compared to Huron and Michigan lakes, the Lake
Superior alone comprises of 82,414 km². However, Huron still has the longest coastline of 6,157
km.
• The world's smallest geological ocean, the Caspian Sea having a surface area of 394,299 km²
which is greater than the six largest freshwater lakes combined, and it's frequently cited as the
world's largest lake.
• The deepest lake is Lake Baikal in Siberia, with a depth of 1,637 m and the mean depth is
also the greatest in the world (749 m). It is also the world's largest lake by volume (23,600 km³,
though smaller than the Caspian Sea at 78,200 km³), and the second longest (about 630 km
from tip to tip).
• The longest lake is Lake Tanganyika, with a length of about 660 km (measured along the
lake's center line). It is also the second largest by volume and second deepest (1,470 m) in the
world, after lake Baikal.
Note : The world's oldest lake is Lake Baikal, followed by Lake Tanganyika (Tanzania).
• The world's highest lake is the Crater lake of Ojos del Salado, located at 6,390 m (20,965 ft).
The Lhagba pool in Tibet at 6,368 m (20,892 ft) comes second.
• The highest large freshwater lake in the world is lake Manasarovar in Tibet an autonomous
region of China.
• The world's highest commercially navigable lake is Lake Titicaca in Peru and Bolivia located
at 3,812 m (12,507 ft) above sea level. It is also the largest freshwater (and second largest
overall) lake in South America.
• The world's lowest lake is the Dead Sea, bordering Israel and Jordan located at 418 m (1,371

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ft) below sea level. It is also one of the lakes with highest salt concentration.
• Lake Huron has the longest lake coastline in the world of about 2980 km, excluding the
coastline of its many inner islands.
• The largest island in a freshwater lake is Manitoulin Island in Lake Huron, with a surface
area of 2,766 km².

Unit 3- Nature of Inland water environment

3.1.1. Nature of Inland water environment
Nature of lake environment
Lake basins with steep incline of bottom at the shore regions have margins which are less
subject to changes. In lakes with bordering low-lying swamp, bog or marsh areas, the margin
shifts with elevation.
High and low water marks
The high water marks can usually be identified by ridges of debris and of certain bottom
materials. Low water marks are easily recognized from the positions of the more prominent
animal an plant zones of shallow waters.
Shore dynamics
Water is restless and during calm period some form of motion varies from relatively gentle to
violent. The inland lakes the principal form of water movement produces shore changes. In
lakes, particularly those of glacial origin, the shore line is much regular and simplified. The
modification of original shore line has been accompanied by two main processes viz, shore
cutting and shore building.
The shore cutting take place by the force of waves when the crest of oncoming wave is more or
less to the shore line and the final plunge of a wave lashes against the opposing land loosening
a certain amount of it. If the shore is composed of glacial drift or of soft materials, it will yield to
continuously bombarding of waves. However, in regions of rocky areas the shore cutting is
slowed down but the erosion is facilitated by rock fragments which are picked by the waves.
On the other hand the shore building results from several processes producing additions to the
original lake margins. Exposed sandy beaches form a beach building during summer by way of
waves coming on to the gently sloping where depth is less than the wave depth and thereby
pushing and carrying ahead some of the sand. Under favorable conditions, the end result is
substantially increased breadth of beach (above water level).
Morphometry 
It can be defined as the study that deals with measurement of significant morphological features
of the basin of a body of water and its included water mass is known as morphometry. Many
fundamental ecological relations are directly dependent upon structural relations of water it is
necessary to make measurements of various morphological features. Following general and
morphometric information should be generated before studying structural and functional
attributes of the system.

Before taking up the morphological studies of a lake, general information regarding type,
historical background, location and general physiography should be collected. 
• Type : The type of body of water viz, lake, pond, marsh, swamp, well, spring, stream, river,
estuary, should be noted. 
• Location : The locality, latitude, longitude and altitude at which the study area is situated

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should be noted from authentic maps. 

• Historical background : Collect the information pertaining to geological history of the basin
and surroundings of natural waters. For artificial bodies the construction or excavation details
are of importance. 
• General physiography : Salient physiographical features related to basin, bank and
catchment area of the body of water should be noted. This includes the features of bed-rock,
coarse gravel, fine gravel, debris, mud, marl, peat, sand, silt, clay, marshy, swampy etc.
The following morphometric parameters are of great importance.
1. Area: The surface area of water-spread can be calculated from a shore – line map of the
body of water.
2. Bathymetry: A bathymetric or contour map is one which denotes the depth at different points
in the body of water.
3. Maximum length: It is the length of line connecting two most remote extremities of the body
of the water.
4. Maximum effective length: It is the length of line connecting two most remote extremities of
the body of water along which wind and wave actions occur without any kind of interruption.
Maximum length and maximum effective length may be the same in most cases.
5. Maximum width: It is the length of straight line connecting most remote transverse
extremities of a body of water.
6. Maximum effective width: It is the length of straight line connecting more remote transverse
extremities of a body of water along which wind and wave actions occur without any kind of land
interruption. 
7. Mean width or Mean breadth ( b ) : It is equal to the area divided by maximum length ( b ) =
a/l
8. Depth: It is the vertical distance between the surface and the underlying bottom.
9. Maximum depth: It is the depth measured at the deepest point.
10. Mean depth: It is calculated by dividing the volume of the body of water by its surface area
( z ) = v/a = Volume / area.
11. Outline map: Representing the outline structure of a lake in a plane surface is called outline
map.
12. Topographical map: Representing various layers of lake basin on a flat surface is called
topographical map.
13. Bathymetric map: Map representing the structure and lake basin is called bathymetric map.
This can be derived from outline map and topographical map.
14. Relative depth (Zr)a2 in ha: It is the ratio of maximum depth in meters to the square root
of area in hectares. Zr = dm /  
15. Shore line: Shore line may be measured on a map by using an instrument called rotometer.
Area of the surface and each depth contour is measured by a digitizer or a polar planimeter.

3.2.1. Physical Characteristics

Pressure
Water is a heavy substance. Pure water weighs 62.4 lb (pounds) per cubic feet at 4°C. This is a
direct result of density. Since, density changes with differences in temperature, compression,
substances in solution and substances in suspension; the weight of a cubic foot of natural water

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is not always the same. The pressure at any subsurface position is the weight of the
superimposed column of water plus the atmospheric pressure at the surface. As depth
increases, the pressure in water is rapidly become great, so that ultimately a crushing effect is
imposed upon objects submerged to considerable depths. This collapse under pressure is
called implosion. The pressure change in lakes and reservoirs are very small than compared to
sea. In lake, having maximum depth of 100 ft., the pressure in the deepest region is about 58 lb.
per sq. in. (4 atmospheres).
Compressibility
Water is virtually incompressible. The coefficient of compressibility for each atmosphere of
pressure is usually given as 52.5 x 10- 6 at 0°C for pressures of 1 to 25 atmospheres. Lake
Superior waters, suddenly rendered absolutely incompressible, would rise in level about 23 cm
and an ordinary inland lake with the maximum depth of 100 ft. under the same circumstances,
would rise about 0.25 mm. Since, increasing pressure compresses the water, thereby
increasing its density to the same slight extent, objects sink in water of uniform temperature at
essentially the same rate at all levels.
Density
Some of the most remarkable phenomena in Limnology are dependent upon density relations in
water. The density of water depends on the quantity of dissolved substances, the temperature
and the pressure. With increasing amounts of dissolved solids the density increases in a roughly
linear fashion. The quantity of dissolved solids for inland waters is usually below 1 g / l, except,
for mineral waters (springs) inland salt water bodies, and water bodies subjected to marine
influence. The density difference due to chemical factors is not more than 0.85 g /l and the
density differences occurring in different zones of the same water body are usually an order of
magnitude less than this. 
i) Variations due to pressure
Water at the surface, subject to a pressure of only 1 atmosphere, is considered as having a
density of unity (1.0); at a pressure of 10 atmospheres, the density is about 1.0005; at 20
atmospheres, the density is about 1.001; and at 30 atmospheres, it is about 1.0015.
ii) Variations due to Temperature
Pure water forms ice at 0°C, and steam at 100°C, but there is change in the density of the liquid
due to temperature. Water possesses the unique quality of having its maximum density at 4°C
and it becomes less dense when the temperature decreases from 4°C to freezing point. Density
of water will be less during summer and it will be high during winter. Sea water becomes heavier
at 0°C. The temperature of maximum density of sea water is 0°C, where as for fresh water it is
4°C.

iii) Changes due to dissolved substances

The total amount of dissolved substances in freshwater is less than that in sea water. Such
substances usually increase the density of water, the amount of increase depending upon the
concentration of dissolved materials and their specific gravity. Evaporation increases the density
by concentrating the dissolved materials and the dilution reduces the density.
iv) Changes due to substances in suspension
All waters contain some suspended particulate matter. The quantity and quality of these
substances vary greatly in different waters and at different times. Silt and certain other materials
are heavier than water and thus increase its weight and other material may have a specific

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gravity similar to that of water and cause no significant change in weight. Density currents and
related phenomena may be caused by substances in suspension. 

3.3.1. Physical characteristics

Mobility (Viscosity)
Water is an exceedingly mobile liquid. Nevertheless, it has internal friction (viscosity). This
viscosity varies with the temperature. Water is distinctly more mobile at ordinary summer
temperatures than that are just before it freezes. The viscosity changes with temperature. The
response of water to wind of fixed velocity would differ with different temperature of the water.
Pressure does not cause any significant change in viscosity. 

Buoyancy is the direct outcome of density and varies with the same factors. The law of
Archimedes states that the buoyancy of an object is equal to the weight of the water it displaces.
The greater the density, the greater the buoyant force; the denser the water, the floating object
will ride higher in the water. Thus, ship passing from fresh water into sea water rises little higher,
and the same ship with the same load would ride somewhat higher in winter than in summer.
Movement of water
The principal forms of movements of water are waves, currents and seiches.
a) Waves
Waves are mainly produced by wind. They occur on every body of water in forms and
magnitudes depending upon various local conditions, such as area of open water; direction, and
velocity of winds; shape of shore line and relative amounts of deep and shallow water. The
greater the expanse of water over which the wind blows the greater the potential wave height,
wave length, and wave velocity. Stevenson (1934) formulated a formula for computing the
maximum height of wave in small bodies of water as
h = 1/3 √F
h = Maximum height in water
F = Fetch of the wind in km.
In open water two types of waves are formed namely waves of oscillation and waves of
translation.

 i.Waves of oscillation: In this type of wave, the water particle moves up and down but no
horizontal movement of water.
 ii.Waves of Translation: In this type of wave there is definite forward movement of water

Depth of wave action in water is of considerable limnological importance, but information about
this is lacking. It has been claimed that in the sea, wave action may exert an influence to a
depth of 182 m.
b) Currents
Currents in lakes are mainly of three kinds, viz, vertical, horizontal and returning. True vertical
currents seldom occur in inland lakes, but may be present in large waters such as the Great
Lakes. When present in inland lakes, they are the result of some unusual thermal,
morphological, or hydrostatic circumstance and upwelling of water from deep water source.
Horizontal currents (undertow currents) are common in lakes. They are usually produced by
wind and often modified by the shape of shore line and form of the basin. The ratio of wind
velocity to water movement diminishes as the wind velocity increases. Also, water velocity
diminishes with the increase in depth.

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Returning currents are formed when water is piled up on an exposed shore as a result of an
onshore wind. Such action raises the water level at the position, and, as a result, the excess
water may return underneath along the bottom. The magnitude and duration of such currents
depend upon the velocity and duration of the wind. Steady vigorous, onshore winds may set up
return currents which extend to the opposite side of the lake.
c) Tides
In inland lakes, tides are almost imperceptible, even in the Great lakes. Lake Michigan is said to
have a tide of about 5 cm. This virtually means that tides in freshwaters are so far as known is
negligible phenomena in Limnology.
d) Seiches
In lakes and along the sea coasts, oscillations of the water level occur under certain
circumstances which are called seiches (pronounced as Saches). A seiche consists of a local,
periodic rise and fall of the water level. It is an example of standing wave in which the water
particles do not travel in circular orbits but the advance and return of the particle are in the same
path. Any influence which produces a temporary, local depression or elevation of water level
may produce a seiche.

3.3.2.Seiches
Most commonly produced seiches in lakes are due to :
1. Winds, temporarily strong, which pile up water on the exposed margin of the lake
2. Sudden change in barometric pressure over a portion of the lake area
3. Earthquakes
4. Land slides
5. Sudden, very heavy rainfall at one end of lake. 
The amplitude depending upon the dimensions of the lake and the intensity of the initial cause
may vary from a fraction of a centimeter in small lakes to 1 m or more in large ones. In lake
Geneva, Switzerland it is reported that the amplitude of a seiche may reach about 2 m. 
Forel (1895) used the following formula for computing the period of oscillation of a seiche in a
lake whose basin has definite regularity of bottom.
t = l/√gh 
where, t = time of one half oscillation in sec
l = length of axis of seiches in meters
g = acceleration of gravity (9,809 m/sec2)
h = depth of water in meters
More complicated formulas were worked out for lakes having irregular basins. Whipple (1927)
presents the following formula.
t =2 l / 3,600√dg
where, t = time of oscillations in hours 
l = length of lake (or length of axis of seiche) in feet
d = mean depth in feet along axis of seiche
g = acceleration of gravity (32.66 ft/sec2) 
Seiche condition in lake Erie, the calculated period is 14.4 hr.
Forms of Seiches
Forel (1895) showed that seiches are of different forms as follows
•Longitudinal seiches - whose axis corresponds with the direction of the long axis of the lake. 

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•Transverse seiches - whose axis lies in the direction of one of the shorter axes of the lake. 
Both longitudinal and transverse seiches are of three different forms :
a) Uninodal - having one node 

b) Binodal – having tow nodes

c) Dicrotic seiche – having two beats (show as two peaks on a limnograph) due to interference
of unimodal and bimodal seiches.

d) Plurinodal – having several nodes

Lesser forms of water motion are sometimes called seiches as for example the short–period,
back and forth flow of water though narrow channels in certain localities in very large lakes and
the subsurface seiches, a type which has been postulated as the cause of certain submerged
currents in lake Erie.

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Subsurface waves, sometimes produced in large bodies of water, occur where subsurface water
is denser than the overlying water. A strong localized wind starts an impulse (wave) in the
underlying layer of water which moves forward in the direction of the wind. As this wave moves
along the warmer lighter water just passes over the crest of the wave but in the opposite
direction, thus producing a surface current opposite to the direction of wind.

Subsurface seiches usually arise from a temporary displacement of the thermocline by the
weight of piled up surface water on one side of a lake due to strong wind action. 

3.4.1. Physical characteristics- Surface film, Temperature

Surface film
When water is exposed to air, it acts as if it were encased within an extremely thin elastic,
surface membrane. This boundary is commonly known as the surface film and is interpreted as
a manifestation of unbalanced molecular action. However, at surface film, there is a surface
tension due to unbalanced attractions between water molecules at surface on one side only and
upward attraction is lacking because there are no water molecules above them.
Surface tension is maximum in pure water than in any other liquid except mercury. Surface film
provides support for organisms and miscellaneous particulate material, upper as well as under
surface of surface film offers mechanized support.
Plants are pleuston whereas animals which are associated with the surface film are termed as
neuston (minute and big).
Effects of surface film
a)Beneficial effects are (i) mechanical support and (ii) respiration mainly air breathing aquatic
insects.
b)Harmful effects are (i) reduction of light penetration thereby it will have effects on
photosynthesis and (ii) traps the minute organisms thereby fall easy prey to big animals.
Temperature
Temperature is one of the most important factors in an aquatic environment. In fact, it is
possible that no other single factor has so many profound influences and so many direct and
indirect effects.
Diurnal and seasonal variations are very much common in freshwater environments than in
marine environment. A diurnal variation range of 4.8 to 5°C has been recorded in a tropical
pond with an average depth of 3.0 m. In shallow water bodies within an average depth of 1.5 m,
the lowest night temperature was 26.6°C. The highest day time temperature was 32°C with a
variation of 5.4°C. In flowing water bodies like streams and rivers there is no such wide
fluctuations in temperature.
Lentic waters of lakes and ponds undergo thermal stratification phenomenon according to

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seasons. Thermal stratification has been reported most frequently in the lakes of tropical
countries such as Java, Sumatra and India.
According to temperature relations lakes have been classified into three types
1)Tropical lakes : In which surface temperature are always above 4°C. 
2)Temperate lakes : In which surface temperature vary above and below 4°C. 
3)Polar lakes : In which surface temperature never goes above 4°C. 
Decrease in temperature cause reduction in metabolism resulting in lower rate of food
consumption. Extreme higher or lower temperature has lethal effects on the aquatic organisms.
Fluctuation in temperature of water regulates the breeding periods, gonodal activation and
thermal induced migration. On the basis of their ability to tolerate thermal variations, most fresh
water organisms are classified into stenotherm and eurytherm. Stenothermic are the organisms
with a narrow range of temperature tolerance while the eurythermic are those organisms with a
wide range of temperature tolerance.
Source of heat for evaporation
a)Sun
b)Water
c)Surroundings
Inland waters are subjected to very extreme variation of temperature due to small expanse and
shallow areas and get heated rapidly during day and are cooled at night.
Rate of evaporation is determined by several factors such as
a)Temperature
b)Relative amount of free surface area of the water
c)Vapour pressure
d)Barometric pressure
e)Amount of wind action
f)Quality of water ie. fresh or salt
e) Thermal conductivity
The thermal conductivity of water is very low. Heat coming to a lake from the sun as partially
absorbed and to some extent conducted, but the really effective heat distribution is due to wind
action in agitating the water and to a much more limited extent, to convection currents.
f) Convection
Convection is the process of the transfer of heat by the movement of heated particles
themselves. For eg, when water in a beaker is heated by a flame placed below it, that portion of
water first heated, expand and rise while the upper, colder, denser portion sink. If the heat
supply continues for some time, there are thus set up ascending and descending currents by
means of which heat is carried all through the total water mass. This form of heat distribution is
known as convection. Most forms of artificial heating of water are of this type.

Convection does occur under the following conditions:

Cooling and sinking of surface water as when the sun sets and under conditions of falling air
temperature
a) Entry of colder water from a tributary
b) Cooling of surface water with the passage of autumn into winter
c) Alterations of winds and calm conditions

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d) Entry of cooler subterranean water at a high level in the basin

e) Advent of rain in temperate region
f) Cooling of the surface water by evaporation

All the plants and animals have an adaptation to certain range of temperature ie. - 200°C to the
boiling point of +100°C. Some can withstand very low temperature for a short duration in an
active state and some blue green algae and bacteria living in hot spring (mineral) condition exist
at temperature up to 90°C, however they reproduce at a slightly lower temperature.

3.4.2. a) Thermal stratification

In tropical lake, heat intake at the surface leads to the formation of a vertical temperature
gradient, within which the thermal resistance become too great for the existing winds to continue
mixing the whole water masses. The upper warmer layer is called epilimnion and the lower
cooler layer is called hypolimnion. In between the two distinct portions, a layer called
thermocline.
Summer stratification
In summer, there are three distinct layers are called epilimnion (upper layer), a bottom layer
called hypolimnion and the middle layer called thermocline or metalimnion.
Epilimnion
a) It is upper layer of water.
b) It is warmer layer.
c) The temperature of this layer fluctuates with the temperature of the atmosphere. It will be
about 27°C to 21°C. 
Hypolimnion
a) It is the bottom layer of water.
b) At this layer, water will be cool.
c) The temp is between 5°C and 7°C. 
d) It is a stagnant column of water. 
Thermocline (metalimnion)
a) It is the middle layer.
b) The temperature is in between the temp of the upper layer and that of the lower layer.
c) It is characterized by a gradation of temperature from top to bottom.
d) It is also called transition zone.
In deeper lakes, a seasonal, thermal phenomenon occur which is so profound and so far
reaching in its influence that it forms, directly and indirectly the substructure upon which the
whole biological framework rests, particularly in the temperature zone. Therefore, a clear
understanding of the salient features of thermal stratification is a necessity.

Thermal relations during spring

Uniform temperature of 4°C prevails throughout the water column of the lake. Wind depresses
water at windward side and drives towards leeward side (towards the sheltered side), sinks at
this end and moves at the bottom. This results in through mixing which is known as isothermic
or homothermic condition.
During summer
As spring advances warmer winds and sun’s radiation increases surface water temperature.

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Water expands above 4°C and thus water at the surface is lighter than underlying colder water.
Upper layers become more warm and lighter and no mixing can takes place. Wind drives water
towards leeward and it sinks at that side which will sink down but not reaching the bottom of the
lake but will be stopped at some intermediary level above cooler (colder) bottom water
(Hypolimnion). 
Currents in the upper lake will induce a counter current which is of a lesser magnitude in the
bottom lake. At this depth, the current direction will be towards the opposite side of the lake ie,
windward side from leeward to wind ward, sinks at this end and returns as the counter current at
this region of lake. Thus two distinct layers are seen at this time in the lake. Between these two
layers, temperature drops suddenly, upper layer in contact with the warmer waters of upper lake
which is mixing by warmer winds and conduction.
On the other hand, the lower layers of this region is in contact with the layer which is yet to gain
heat through conduction and other processes which are themselves slow process. This
separating zone between upper lake and bottom of lake is called as Thermocline region. It is
defined as a region wherein the temperature drops by more than 1°C per meter of depth. The
term of thermocline was proposed by Birge (1897). Thus, epilimnion / upper lake is above
thermocline and bottom lake / hypolimnion is below region of thermocline.
During fall (autumn)
Cold wind blow over the lake surface which cools surface water which become denser at -4°C.
These denser waters sink through lighter warmer waters to a level where it meets the waters of
similar density ie. first it will be at thermocline. Thus epilimnion gradually cools and on the other
hand the hypolimnion will maintain the same temperature. A stage will be reached when there
will be no thermocline region, water freely mixes. This mixing is called fall overturn. Mixing
continuous till the temperature throughout will be at 4°C.
During winter
Cooling below 4°C will make water lighter and thus the surface waters are lighter than the
warmer but denser subsurface water. This water floats and no sinking, cooling continuous at
surface till ice is formed at 0°C. Once ice is formed at the surface wind has no effect as far as
mixing is concerned a period of stagnation sets in.

During spring
With the onset of spring, warmer sun rays and wind melt the ice cover. Now colder but lighter
water will be above warmer but denser water below. Once it attains a temperature of 4°C, it
sinks down and reaches a level of 1°C which being lighter ascends up and in turn warms up.
Thus the layer of denser water increases until the whole lake is uniformly of a same
temperature. Mixing takes place now by spring winds and this is called as spring overturn.

3.5.1. Physical characteristics- Light, Colour, Turbidity

Light
Light influences freshwater ecosystems greatly. Fresh waters contain more of suspended
materials. These suspended materials obstruct the light that penetration reaches the water. The
degree of such obstruction of light influences the productivity of the freshwater ecosystem. A
shallow lake receives light to its very bottom resulting in an abundant growth of vegetation both
phytoplankton and rooted vascular plants. Light affect the orientation and changes in position of
attached species and their nature of growth and it also causes the diurnal migration of
planktonic organisms. The factors affecting the light penetration in natural waters are the

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intensity at the surface, angle of contact of light with surface, differences in latitude, seasonal
differences, diurnal differences and suspended materials.
The light intensity at which oxygen production by photosynthesis and oxygen consumption by
the respiration of the plants concerned are equal is known as the compensation point, and the
depth at which the compensation point occurs is called the compensation depth.
Light exerts a great influence on many biological process of water. Most important future of
water is its transparency. This fluctuates in different seasons and water bodies such as flooding
livers, mountain streams etc. The source of light on the earth - a) Sun and b) Moon
Electromagnetic spectrum emitted by Sun (a) short gama rays (0.0001 mm) to (b) long Hertizan
waves (several km long). The Hertizan waves are the electromagnetic waves used in radio and
it is pronounced as Hertz.
Intensity of light is the number of quanta passing through on a unit area, ie, light energy and the
unit of expression of light intensity is ‘Lux’
Wave length is the measure of light colour
nm = nanometer = mille micron 10-9
AO = 1/ton billionth of a meter
nm = 1/billionth of a meter or 10AO
Intense radiation is restricted to 300 to 1300 nm. Peak radiation distribution is in the blue green
range.
Wave length heating water is 0.1 to 770 nm (infra red spectrum).
In a year the amount of radiant energy that reaches earth from the sun is 1.3x1021 k cal
Visible wave length/light : 400 to 770 nm; Ultraviolet light >286 to 400 nm
Light penetration in natural waters is affected by
a) Dissolved substances
b) Suspended substances
c) Planktonic organisms
d) Geographical features (latitude and longitude etc)
e) Meteorological conditions
f) Angle of light etc.
Methods for estimation
a) Secchi’s disc
Secchi (1865), an Italian professor employed a metallic disc for measuring the transparency of
waters of Mediterranean sea. It considered in lowering into the water a white metallic disc of 20
cm in diameter, on a graduated rope, noting / recording the depth at which the disc disappeared
then lifting the disc and noting the depth at which it reappeared. The average of these two
readings was considered the limit of viability or Secchi disc depth. This method was used
subsequently by many investigators. Whipple modified this method by dividing the disc into four
quadrants and paintings them in such a way that two of the quadrants which were directly
opposite to each other, black and intervening ones white. He also increased the efficiency of the
method by viewing the disc, as it sank in the water through a water telescope held under the
sun shade.
This method is not actual measure of light penetration, but instead merely a useful rough index
of visibility when used under standard conditions. They are (a) Clear sky (b) Sun above the
head (preferably) (c) Shaded or protected side of the boat (d)) Under a sun shade. This method
has come into a wide use as a means of comparing different waters.
Factors influencing the light penetration
1) Intensity of light at surface
This varies (a) degree of clarity of sky (b) presence of fog, dust, smoke etc and (c) time of the
day/season of the year.
2) Angle of contact with surface
Light in contact with surface part of it is reflected rest enters water and becomes refracted.

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Penetration depends on angle of contact and maximum penetration when sun is at zenith.
3) Different in latitude
More remote the water mass is from equator, greater will be the departure of sun’s rays from
vertical and hence penetration varies.
4) Seasonal differences
Closely associated with latitude are the seasonal changes in the position of the sun. Only
locations at or between 23° 28i N and 23° 28i S (Tropic of cancer and Tropic of Capricorn) ever
have a vertical sum. Beyond this zone, north or south not only do locations have on regular sun
but the angle changes progressively with change of seasons.
5) Diurnal difference
Angle of light in contact with water is ever changing during day, reaches zenith at noon.
6) Dissolved materials
One of the important factors is absorbance of light which varies with chemical substances such
as (a) chloride of Ca and Mg affect light penetration ie. Diminishes, (b) Traces of NH3 proteins,
nitrate, carbohydrates etc reduces the light penetration with respect to ultraviolet rays. 
7) Suspended materials
Silt, clay etc. are effectively screen light and also the penetrations of light reduce by
phytoplankton and zooplankton.
Penetration of light in pure water
When light penetrates or enters into pure water (a) certain portion of light is absorbed and (b)
some of it is scattered in the form of deflection in all directions.
Absorption is selective in which certain wave lengths are absorbed more quickly than others.
Penetration of light in natural waters
Every quantitative determination records were only in marine waters probably because of more
clarity. Here photographic plate method used by Forel (1865) in lake Geneva at about 200 m.
3.5.2. Physical characteristics- Colour and Turbidity
Colour
Pure water bodies appear nearly black as they absorb all light components of the spectrum. The
lake water containing suspended materials is seen blue in colour due to the scattering of light by
water molecules. Natural waters differ greatly in colour, depending upon the materials dissolved
and suspended in it. 
It is a common misconception that in large water bodies, such as the oceans, the water color is
blue due to the reflections from the sky on its surface. Reflection of light off the surface of water
only contributes significantly when the water surface is extremely still, ie, mirror like, and the
angle of incidence is high, as water's reflectivity rapidly approaches near total reflection under
these circumstances. Some constituents of sea water can influence the shade of blue of the
ocean and hence it can look greener or bluer in different areas.
Scattering from suspended particles also plays an important role in the color of lakes and
oceans. A few tens of meters of water will absorb all light, so without scattering, all bodies of
water would appear black. Because, most lakes and oceans contain suspended living matter
and mineral particles as coloured dissolved organic matter (CDOM) and thus the light from
above is reflected upwards. Scattering from suspended particles would normally give a white
color, as with snow, but because the light first passes through many meters and the scattered
light appears blue. In extremely pure water as is found in mountain lakes, where scattering from
white coloured particles is missing, the scattering from water molecules themselves also
contributes a blue color.

Turbidity

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Degree of opaqueness developed in water by means of suspended water is known as turbidity.

Turbidity producing substances may be divided into two groups.
i) Settling suspended matters – those substances which in motionless water, will settle to the
bottom sooner or later.
ii)Non-settling suspended matters - Finely divided solids or those materials whose specific
gravity is less than water which are in permanent.
The settling of particulate materials is by no means at a uniform rate, particularly in deeper lake
having considerable difference in temperature between the surface and the bottom layers.
Effects of materials in suspension
a)Light reduction: Favourable for animals but unfavourable for plants (photosynthesis)
b)Effects of temperature: Turbid waters are warmer than clear waters. Suspended particles
absorb heat more rapidly than water itself and then radiate the heat to the surrounding water,
adding to the heat content of the water.

Unit 4- Chemical characteristics

4.1. Dissolved gases – Oxygen, Carbon dioxide and other dissolved gases
Dissolved gases
No naturally occurring body of water is free of dissolved gases. Their spatial and temporal
distribution is dependent on factors such as precipitation, inflow and outflow, physical factors
like temperature, movement of water and chemical factors such as solution processes,
combination and precipitation of reactions, complex formation etc.
Among the dissolved gases present in water, oxygen and carbon dioxide are direct indicators of
biological activity of water bodies. Gaseous nitrogen only enters the metabolic cycle of a few
specific microorganisms. Hydrogen sulphide and methane occur in small localized amounts due
to bacterial activity under conditions of low redox potential and are incorporated into the material
budget of water bodies by certain bacteria.
The Liebig’s law of minimum states that the yield is dependent on whatever growth factor is at a
minimum in proportion to all the other similar factors. 
Solubility of Gases in water
The solubility of gases in water decreases with increasing temperature and decrease of
pressure. When a gas comes in contact with water, it dissolves in it until a state of equilibrium is
reached in which the solution and the emission of the gas are balanced. Total solubility of gas is
expressed by Henry’s law. The concentration of a saturated solution of gas is proportional to the
pressure at which the gas is supplied.
Condition affecting the solubility of gases in water
Solubility of gases differs widely even when their pressures are equal. It is therefore necessary
to find out the solubility constants.
Henry’s law is stated as :
C= K p
Where, C = Concentration of gas in solution
p = Partial pressure of gas
K= Constant of solubility
The following general conditions affect the solubility of a gas:
i. Rise in temperature reduces solubility

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ii. Increasing concentration of dissolved salts diminishes solubility

iii. Rate of solubility is greater when the gases are dry than when they contain water vapour
iv. Rate of solubility is increased by wave action and other forms of surface water agitation

A. Oxygen
The main sources of dissolved oxygen in water are:
i) The atmosphere and
ii) By photosynthetic activity of aquatic plants
Atmospheric oxygen enters the aquatic system:
a) By direct diffusion at the surface and
b) Through various forms of surface water agitations such as wave action, waterfalls, and
turbulences due to obstructions.
Aquatic chlorophyll bearing plants release oxygen as a byproduct of photosynthesis, which gets
distributed into the different layers of lake water by movements. In most lakes the phytoplankton
contributes the bulk of the oxygen supply because of the huge amounts of chlorophyll of algae
in the epilimnion zone. In shallow waters like ponds and swamps the limnetic photoautotroph
may be overshadowed by littoral macrophytes, attached algae, and the benthic algal mats. In
small rivulets and brooks the periphyton account for most of the production of oxygen.
The main causes of decrease of oxygen in water are:
i. Respiration of animals and plants throughout the day and night and
ii. Decomposition of organic matter – Aerobic bacteria use up of the oxygen of water while
decomposing organic matter. Chemical oxidation of sediments also takes place. Purely
chemical oxidation may also occur, but most of the oxidative processes in aquatic habitats are
probably mediated through bacterial action.
iii. Reduction due to other gases – A gas may be entirely removed from solution by bubbling
another gas through the water in which it is dissolved. In nature, gases like CO2, methane and
hydrogen sulphide often accumulate in large amounts and the excess amounts rise in the form
of bubbles removing the dissolved oxygen.
iv. By physical process – In summer days the heat warms up the epilimnion zone of the lake,
which could account for oxygen depletion of water. The combined effects of all or some of the
above mentioned processes may completely deplete oxygen content of the system.
Diel oxygen changes in freshwaters
The concentration of oxygen in an aquatic environment is a function of biological processes
such as photosynthesis and respiration and physical processes such as water movement and
temperature. Diel variations occur in both day and night hours. Estimates of diel production can
be made in natural waters by considering night as the dark bottle and day as the clear bottle.
The increase in oxygen from dawn to dusk reflects net primary productivity. The decrease from
dusk until dawn represents half the diel respiration. Adding the oxygen that disappeared at night
to the day time gain gives a sum that is daily gross primary productivity. 

B. Carbon dioxide
i) Sources of carbon dioxide in freshwater
The atmospheric carbon dioxide mixes with the water when it comes in contact with the water
surface, as it possesses the highest solubility in water. As the partial pressure of carbon dioxide

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in air is low, the amount which remains in solution in water at a given temperature is also low.
1. Rainwater and inflowing ground water
Rainwater is charged with 0.55 to 0.60 mg/I CO2 as it falls towards earth. Water trickling through
organic soil may become further charged with CO2.
2. Byproduct of Decomposing Organic Matter (DOM)
Carbon dioxide is added to the water as a byproduct of decomposing organic matter which is a
common phenomenon in natural waters. Large quantities of the gas are produced in this way. It
is found that carbon dioxide is the second largest decomposition product, constituting 3 to 30
per cent of the total gas evolved.
3. Respiration of Animals and Plants
Respiratory processes produce and release carbon dioxide into the water. The quantities so
added are governed by the magnitude of aquatic flora and fauna, the relative size of the
individual organism and those factors which determine the rate of respiration.
ii) Reduction of carbon dioxide in freshwaters
The principal processes which tend to reduce the carbon dioxide supply are;
1. Photosynthesis of aquatic plants
Consumption of free CO2 in photosynthesis depends upon amount of green plants which the
water supports, duration of effective day light, transparency of water and the time of year.
Marl forming organisms
The following groups of aquatic organisms are known to form marl (=Crumble : large deposits of
calcium and magnesium carbonate) in water bodies; aquatic flowering plants like Potamogeton,
Ceratophyllum, Nymphaea, Vallisneria; many blue-green algae like Rivularia, Lyngbya nana,
Lyngbya martesiana, Colacacia. Centrosphaeria facciolaea; many species of diatoms; mollusks
which form calcareous shells; insects like Diptera larvae; the cray fishes and lime-forming
bacteria. All these organisms function in the production of the insoluble carbonates which
involves carbon dioxide, calcium and magnesium. Thus the process of lime formation binds up
carbon dioxide supplied from circulation and removes the available calcium and magnesium
from the system.
Agitation of water
Agitation is a very effective method of releasing free carbon dioxide from water. It is evident
from the fact that sometimes when deeper layers of water has large amount of it, the surface
water shows very little carbon dioxide.
Evaporation
Evaporation of waters containing bicarbonates results in the loss of half-bound carbon dioxide
and precipitation of mono carbonate. The form of loss is greatest in shallow water bodies where
evaporation is most effective.
Rise of bubbles from depths
Free carbon dioxide often accumulates in decomposing bottom deposit in such quantities that at
frequent intervals increasing internal pressure of gas exceeds the external pressure and the
excess gas rises in the form of masses of bubbles to the surface and is lost into the air.

Other dissolved gases

i) Methane
Methane, sometimes called marsh gas, is one of the products of decomposing organic matter at

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the bottoms of marshes, ponds, rice field and lakes. The methane bacteria are obligate
anaerobes. They decompose organic compounds with the production of methane (CH4) through
reduction of either organic or carbonate carbon. Conditions favorable for production of methane
appear at about the time the dissolved oxygen content is exhausted. This is because methane
(CH4), a compound of carbon and hydrogen burns in oxygen forming oxides of carbon and
hydrogen ie, carbon dioxide and water.

It has been found that large quantities of methane are produced in marshes and eutrophicated
lakes during summer time.
ii) Hydrogen Sulphide
Hydrogen sulphide dissolves very rapidly in water and is thus not dissipated like methane. The
bottom water of stratified eutrophic lakes may contain appreciable quantities of the very soluble
gas H2S. This is especially marked in lakes of regions of high edaphic sulfate. The reduction of
sulfate to sulfide is a phenomenon largely associated with anaerobic sediments. H2S is
poisonous to aerobic organisms because it inactivates the enzyme cytochrome oxidase. 
iii) Nitrogen
Nitrogen has a low solubility in water. It is such an inert gas that the quantities which occur in
lake water are not changed by the chemical and biological processes. The atmosphere usually
supplies the greater amounts of nitrogen found in water. The minimum amount occurs in winter,
since it is more soluble in cool water. 
iv) Ammonia
Ammonia occurs in small amounts in unmodified natural waters. It is exceedingly soluble, 1
volume of water dissolving 1,300 volume of ammonia at 0° C. In lakes, it is the result of the
decomposition of organic matter at the bottom. In summer, free ammonia ordinarily increases
with depth.
v) Sulphur dioxide
Traces of sulphur dioxide may occur in natural waters.
vi) Hydrogen
Liberation of hydrogen in the anaerobic decomposition of lake bottom deposits seems likely.
But, the amount so formed is small.
vii) Carbon Monoxide
Carbon monoxide may occur in the bottom of the hypolimnion in small amount.

4.2.1. Dissolved Solids and Dissolved Organic Matter

All waters in nature contain dissolved solids .Water is the universal solvent dissolving more
different materials than any other liquid. Natural waters come in contact with soluble substances
in many ways such as mere contact with its own basin, erosion at shore line, wind blown
materials, inflow of surface waters, inflow of seepage and other forms of subterranean waters
and decay of aquatic organisms. Rain water contains 30 to 40 ppm of dissolved solids.
Solubility of solids in water
Salts are composed of ions which in the solid form are held together by ionic forces. The strong

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ionization of the salts leads to the formation of hydrates with water in which the water acts as a
dipole to which the ions are attached. The solubility of solid substances is strongly dependent
on the pH and the redox potential in the water. It usually increases with temperature and is
largely independent of pressure. Most substances dissolve either in the molecular form or
dissociated into ions. Some important constituents such as humic acids, salicilic acid and ferric
oxyhydrate are dispersed in colloidal form.
Major ions in freshwaters
The major ion contents vary in different fresh waters due to five factors, which are climate,
geography, topography, biotic activity and time. These are not completely independent and they
interact. 
Carbonate is the principal anion in most fresh-waters. Generally carbonate occurs as
bicarbonate ion with calcium in water. Bicarbonate ion is customarily expressed as
CO3 because evaporation of a known amount of calcium bicarbonate solution leaves only the
carbonate of calcium to be weighed. During evaporation, gaseous CO2 and water are lost, from
bicarbonate ions, converting them to a lesser weight of carbonate.

Evaporation

Alkalinity is usually a measure of carbonates. There are various compounds of carbonates with
calcium, such as calcite or aragonite which have the same chemical formula (CaCO3), but are
crystallized differently. Aragonite precipitates from thermal waters and is contained especially in
the shells of freshwater mollusks. Magnetite, the carbonates of magnesium (MgCO3) and
dolomite, a double carbonate of calcium and magnesium, Ca Mg (CO3)2 are also relatively
common. Carbonates of barium (BaCO3) and strontium (SrCO3) also occur. CaCO3 is insoluble
except in the presence of acid. With carbonic acid, it becomes Ca (HCO3)2. Because of this, it
seems reasonable to express alkalinity titration in terms of bicarbonate ions, but on the other
hand, Ca(HCO3)2 is very unstable and when water is evaporated to determine its contained
dissolved salts, the bicarbonate of calcium is destroyed and only carbonate remains.
Dissolved inorganic solids
i) Nitrogen compounds
Nitrogen occurs in natural waters in the form of numerous compounds, in inorganic form as
nitrate, nitrite and ammonium and in organic form as intermediate stages of microbial protein
decomposition. The most important inorganic nitrogen compounds in water are nitrate and
ammonia. Natural waters contain some ammonium salts. Ammonium carbonate is probably the
common form.
iii) Phosphorus compounds 
Free phosphorus does not occur in nature, but in the form of phosphates it is abundant.
Inorganic phosphorus compounds usually occur in dissolved form only in small amounts in
natural waters, often only as traces. Total phosphorus in lake water includes two components.
One is soluble phosphorus which is the phosphate form and another one is organic phosphorus
which is contained in plankton organisms and other organic matter in the water. As an essential
nutrient for primary producers, phosphorus thus acts more often than nitrogen as the growth

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limiting factor. The natural inorganic phosphate content originate from atmospheric precipitation
as well as from various phosphate containing rocks especially apatite, which are flushed into the
lake by tributary streams. In lakes and flowing waters three phosphate fractions occur
concurrently : soluble inorganic phosphate as orthophosphate (PO4) and polyphosphate,
soluble organic phosphate and particulate organic phosphate (organisms or detritus). These
fractions make up to total phosphate content. The losses of phosphorus occur throughout
flowing water which removes both soluble and organic form. It may also occur through removals
of fish, molluscs, water plants and other organisms.
iii) Sulfur compounds
The inorganic sulfur compound occurring predominantly in natural waters is sulfate. In this form
sulfur can be absorbed by phytoplankters and other photo-autotrophs. Purely chemical
processes involved in the sulfur budget of natural waters are the oxidation of hydrogen sulfide to
sulfur by molecular oxygen and also the formation of sulfides, especially iron sulfide in the
sediment. The sulfate ion, SO4 is usually second to carbonate as the principal anion is fresh
waters, although chloride sometimes surpasses it. Silica often outranks sulfate, but very little is
ionized. Free or elemental sulfur is inactive at ordinary temperature. This element can combine
with both metals and non-metals to form many compounds. Free sulfur is an important
constituent of protoplasm; it is protein and specifically within those amino acids having sulfhydryl
(SH) bonding; e.g. cystine, cyseine and methionine. 
When sulfur is combined with hydrogen the most reduced state is sulfide (S- -) and the most
important sulfides in limnology are the gas - hydrogen sulfide (H2S) and ferrous sulfide (FeS).
Sulfates combine with hydrogen to form sulfuric acid. With the alkali metals sulfur forms the
most abundant form in lakes and streams.
Atmospheric sources of sulfate have increased with man’s industrial activities. Man now
contributes about ten times more SO2 than that the annual contribution from volcanoes. Coal
combustion produces the gas maximum and copper smelting and paper manufacturing add to it.
Through precipitation and runoff water the sulfate level of some fresh water becomes unusually
high showing industrial water pollution.
The most important conversion process for sulfur in lakes can be summarized as follows: sulfate
is reduced by the desulphuricans to H2S and sulfides which are deposited in the sediment.
Hydrogen sulfide is also formed by the microbial decomposition of proteins which is oxidized by
Thiobacteria, Chromatiaceae and Chlorobiaceae via molecular sulfur to sulfate.
iv) Silicon
Silicon does not occur in nature as a free element. Natural waters commonly contain silicon
dioxide in some form of soluble silicate. Silica may also exist in certain waters, particularly in
rivers, in colloidal form. River waters are relatively rich in silica. Silica is the second most
abundant element in the lithosphere. Its main source in fresh water and sea water is weathering
of the feldspar rocks. In inland waters it ranges from 0.1 ppm. Solubility of silica increases with
the rise in temperature. Dissolved silica remains as H2SiO4. Silica is an essential nutrient for
diatoms as they build up their frustules with this material. The decay of silica is slower than
organic compounds and thus many diatoms frustules may be buried and lost to the lake
sediments. 
Other Elements
Certain elements such as calcium, magnesium, manganese, iron, sodium, potassium, sulphur,

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copper and others constitute elements of chemical compound dissolved in the water.
i) Calcium
The predominant compound of calcium is CaCO3, which is very less soluble in water but in the
presence of carbonic acid it is represented abundantly as the soluble Ca (HCO3)2. Thus there is
an inseparable relationship between carbonic acid, CO2, pH and the anion CO3-in water.
The earth’s crust contains an ample store of calcium as a constituent of certain silicates.
Anorthite (CaAl2Si2O8) is a common member of the feldspar group of silicates. They are the
most abundant of all minerals and make up 60% of the earth’s coating. A deposit of sedimentary
CaCO3 is changed to soluble bicarbonate by the action of CO2 rich meteoric water (rain water)
which enters aquatic systems. The solubility of CaCO3 depends on CO2 which follow the
reaction: CaCO3 – Ca (HCO3)2. This equilibrium is disturbed when the water emerged and the
pressure on it suddenly released making the escape of CO2. Also removal of CO2 in photo-
synthesis will disrupt the equilibrium. CO2 is assimilated in photosynthesis and CaCO3 gets
precipitated in the form of calcareous incrustations on plants and other submerged objects.
Other Minerals of Calcium
After the silicate and carbonate minerals of calcium, the sulfates rank as its most abundant
store. These are gypsum, (CaSO4, 2H2O) and anhydrite (CaSo4).
ii) Magnesium
Magnesium is usually the second most abundant cat ion in inland waters. Its source is both
silicate and non-silicate minerals of the earth’s crust. Foresterite (Mg2SiO9) in the following
manner:

The magnesium carbonate is called magnetite and a double carbonate is dolomite, Ca Mg

(CO3)2. Epsom salt (Mg SO4.7H2O) is a soft and whitish sulfate of magnesium. It is 150 times
more soluble than gypsum (CaSO4.2H2O). Epsom salt occurs in mineral spring deposits and in
salt sediments of certain lakes.
iii) Sodium
The monovalent alkali metal is very reactive and soluble. When leached from the rocks, its
compounds tend to remain in solution. For this reaction, it is at least the third most abundant
metal in lakes and streams and sometimes it ranks first. Among igneous rocks, the feldspars,
alumino silicates of alkali and alkaline earth metals are the most abundant of all minerals. The
commonest water soluble mineral is halite or simply Na Cl. In arid tracts like Rajasthan where
closed basins hold concentrated waters, there are at least 3 types of sodium lakes:
i) Slatterns, having concentration of sea water with preponderance of Na Cl.
ii) Saline lakes having Na2SO4 in water and
iii) The soda lakes characterized by NaHCO3 and Na2CO3.
They have been termed as alkali water because of their high pH. These soda lakes are
characterized in having luxuriant growth of blue-green algae.
iv) Potassium
Potassium, a close relative of sodium is usually the fourth ranking cat ion in freshwater. In usual
cases it may surpass sodium in certain lakes. It is weathered from various feldspars that have
the formula KAISi3O8 but does not remain in solution. Potassium also tends to form plates of

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mica, which are insoluble and unavailable to aquatic ecosystems. Because of this formation
potassium becomes rarer in water than sodium. In plants, both extracellular and intracellular
fluids contain an excess of K+ over Na+. In animals extracellular Na+ often surpasses
potassium. There is some evidence that highly concentrated water with a pronounce potassium
content are lethal to many aquatic animals, the Na/K ratio being less than ten.
Potash is the name for K2CO3 but he word has been used to refer to KOH or potassium oxide.
The so called potash lakes occupy depressions among the sand in Nebraska in USA.
v) Iron and Manganese
Although iron is one of the most widely distributed elements on the earth, it occurs in natural
waters only in relatively small amounts due to its specific solubility properties. But the ground
water contains large amounts of dissolved iron and manganese. The compound of trivalent
(ferric) iron is almost completely insoluble in water. Thus iron remains in solution only in the
bivalent form, under reducing conditions, and chiefly as the bicarbonate Fe (HCO3)2. The
conditions under which bivalent iron (ferrous) compounds remain in solution are: an oxygen
saturation value of less than 50%, the presence of degradable organic matter, a high level of
free CO2 and a pH of less than 7.5. These conditions are found primarily in groundwater and in
the hypolimnion of lakes.
vi) Chloride
It is an element of the halogen group that includes also fluorine, iodine and bromine. Among
these members, chloride surpasses them in polluted as well as freshwater lakes and streams.
Molecular chloride (Cl2) is a heavy yellow lethal gas, but in natural waters, it is dissociated as
chloride ions, which combine with all common cat ions. It is stored in most freshwater algal cells.
Contamination of water from domestic sewage can be monitored by chloride assays of the
concentrated water bodies. This is because a human and animal excretion contains an average
of 5g Cl- per liter.
Dissolved Organic Matter 
Freshwater contains 0.1 to 50 mg dissolved organic compounds (DOC) per litre. Various free
sugars, amino acids, organic acids, polypeptides and other substances have been reported.
There are probably four sources of these dissolved materials
1. Organic compounds of allochthonous origin
2. Soluble organic material from the decay of aquatic organisms 
3. Extra cellular metabolites excreted by littoral macrophytes
4. Excretion from the fresh water animals.
The organic compounds not only serve directly as source of energy but also are associated with
the nutrient cycle of the ecosystem. Most metals are transported down streams or exist in lake
making complexes with organic materials either being absorbed or occurring as metallic
coatings on detritus. Such organic substances as humic material causing yellow stain in fresh
water comes from the decay of plant material in the soil.
The origin of the dissolved organic compounds in the water is manifold. The losses are due to
photorespiration, secretion of the products of algal photosynthesis and those of higher plants
and also due to the excretions of bacteria. 
An important group of organic substances in water consists of humic substances. They are
polymeric mixers derived mostly from such plant materials as lignin. Cellulose, proteins and fats
humic substances enter the water due to incomplete breakdown of plant residues in the water

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bodies.
They affect the material budget in as much as they enter into complex-formation with heavy
metals (iron and manganese) and consequently prevent their precipitation and ensure their
continued availability to the primary producers. Thus a direct relationship may be observed
between the concentrations of dissolved iron and water soluble humic substances in lakes.
Heavy-metal ions may also become adsorptive bound to particulate humic materials. This
tendency to chemical and adsorptive binding of heavy metals is of great importance for
productivity in natural waters. Calcium may precipitate as calcium humate on contact with humic
acids and be deposited in the sediment.

Humic acids in sediments also form iron-humus-phosphate complexes and hence, phosphate
combination with iron is substantially reduced. The humic materials hold in suspension. The
humic materials hold in suspension large quantities of metallic ions and through their chelating
activity hold on the essential trace metals are delayed in the presence of humic compounds and
thus hold the nutrients for a longer period in water.

Unit 5- Biological relations

5.1 Influence of physical and chemical conditions on living organisms in inland waters 
Shoreline
The greater the length of the shore line the greater the biological productivity. Increased
irregularity of shore line results in
1) greater contact of water with land
2) increased areas of protected bays and covers
3) increased areas of shallow water for growths of rooted vegetation
4) greater diversification of bottom and margin conditions
5) reduction of the amount of exposed, wave-swept shoal and
6) increased opportunity for extensive, close superposition of the photosynthetic zone upon the
decomposition zone. These and other possible results combine in various ways to increase the
production of animals and plants.
Relation of photosynthetic and decomposition zones
The form of basin of a lake determines, among other things, the relative amounts of shallower
waters. Within limits and under strictly comparable conditions, the grater the areas of shallow
water the greater the biological productivity. Their exposed nature usually results in
a. the absence of rooted plants
b. the absence of organic bottom deposits and
c. the absence of any permanent animal population save those whose burrowing habits make
occupancy of position in such a habitat possible.
Nevertheless, a fairly substantial but largely concealed population may be present, although of
all the shallow water faunas it is usually the smallest one.
In contrast to conditions described above, the steeper the slope of the basin, or the greater the
exposure of shoals, or both, the greater the removal of the decomposition zone to the profundal
depths of deeper lakes

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A deeper lake with steep basin slope thus tends to automatically and continuously rob itself of
its stores of organic matter. Lakes of the third order, because of shallow depth and continuous
circulation except during the ice cover period, retain all organic accumulations in available
position, the essential decomposition products either remaining immediately beneath the plant
beds or else being constantly the tribute by the water.
Slope and the deeper decomposition zone
Form of basin also involves the slope of deeper portions of the basin. Upon the nature of this
slope depends, to a large extent, the character of the bottom. The influence of gravity, aided by
water movements in pulling to the lowermost bottom the various materials which settle through,
is much more effective on a declivitous slope. Some basin slopes are so abrupt that very little of
the loose, settling materials can remain on the steep sides. Thus the decomposition zones of
such a lake are restricted to (1) those of the shallow, protected shoals (if any are presently), and
(2) those at the bottoms of the deepest regions, separated by steep sides which maintain little or
no decomposition deposits.

5.2. Productive volume, flotation phenomena and body form adjustments

Productive volume
Form of basin determines the extent of productive volume. By productive volume is meant that
portion of water in which virtually all biological production occurs. In a lake of the third order,
total volume is productive at least during the open season. In lakes of the second order,
productive volume is almost exclusively confined to the epilimnion and the thermocline during
most of the summer stagnation period. During the overturns, the entire lake temporarily
becomes productive volume but the duration of these periods may be too short to be or any
great consequence. Lakes of the first order resemble those of the second order in that they
maintain the productive zone in the upper stratum, usually limited by the presence of a
thermocline. In those lakes having no complete overturn, the productive zone, during open
season, merely varies in volume with those conditions which determine the depth to which
circulation may extend. During prolonged ice cover, lakes of the second and third orders
undergo gradual reduction of productive volume due to encroachment of the underlying
stagnation zone, while lakes of the first order, under these conditions, may undergo less change
in productive volume due to their size and to the presence of the permanent, deeper stagnation
region. Complete ice cover may not occur in lakes of unusual size and depth, even though
located in colder regions. 
Both depth and area of a lake basin combine in innumerable ways to produce a great
heterogeneity of lake forms. Since inland lakes seldom exceed certain area limit, depth is
fundamentally of prime importance in determining productive volume.
The average depth is the factor which determines whether a lake is eutrophic or oligotrophic,
computing average depth as the quotient of volume of the lake over area of the lake (V/A). In
oligotrophic lakes, the volume of the hypolimnion is greater than the volume of the epilimnion,
and that in eutrophic lakes the reverse occurs.
Flotation phenomena
i). Non-motile organisms
In calm water, non-motile plankton organisms depend entirely upon the relation between their
own specific gravity, the density of the water, and the viscosity of the water in maintaining their
vertical position. Non-motile, attached animals, particularly the colonial forms such as the larger

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colonies of fresh-water sponges and certain fresh water Bryozoa (Pectinatella), may develop
forms and masses of body, which could not be maintained in the absence of the buoyant effect
of water, and even the soft bodied Hydra would be helpless without it. 
All sessile animals depend, to some extent at least, upon the buoyancy of the water. Many
higher aquatic plants are dependent upon the supporting effect of the water in order to maintain
their proper form and orientation.
ii) Motile Organisms
Those plankters which possess powers of locomotion vary greatly in the efficiency of their
progression, but some change of position in space is possible due to their own activity. While
such locomotion may be almost negligible when compared with the shifting and transporting
effects of the water, it may nevertheless be vital to the organism in many ways, such as in the
capture of food, and in the change of water in contact with respiratory surfaces.
Locomotion in water, consumes less energy to maintain their position above the bottom, due to
buoyant effect of water. In fact, those organisms whose specific gravity is essentially the same
as the surrounding water expend practically no energy in merely keeping up in the water.
Certain aquatic animals, because of the possession of air stores or other special means, are
distinctly lighter than water and must use a certain amount of energy to keep below the surface
when they need to do so.
Many air breathing, aquatic insects have air stores so located that not only are they lighter than
water but the posterior end is lighter than the anterior, enabling the insect to float at the surface
in the proper respiratory position.
iii) Reduction of specific gravity
Protoplasm alone has a specific gravity which closely approaches that of water, but the various
cell products which occur in animals and plants may combine to produce bodies which are
either heavier or lighter than water. Products which tend to make the body heavier (such as
chitinous exoskeletons of arthropods, bones, shells or various kinds) and those which tend to
make it lighter are often present in the same body so that the specific gravity of the whole
depends upon which of the contrasting materials predominate. The most effective and the most
common of those cell products which reduce specific gravity seem to be the following:
1.Gases originate from various sources (metabolic products, external and internal air stores,
and others) and remain, at least for a time, enclosed within or attached to the body. These gas
accumulations may be of sufficient magnitude to make an otherwise heavy-bodied animal (as,
for example, certain aquatic insects) much lighter than water.
2.Fats and oils are commonly produced and stored within aquatic organisms, notably in the
plankton crustaceans and in the plankton ie Algae. 
3.Gelatinous and mucilaginous secretions of varying amounts are common as matrices and
external envelopes which helps in flotation.
iv) Relations of surface to volume
(a) In different Species. 
Since in relatively compact bodies of organisms, the relations between volume and surface tend
to conform roughly to the well-known mathematical principle that the surface varies as the
square of the dimension while the volume varies as the cube of the dimension. The smaller the
body, the greater will be the relative expanse of surface.
If the body has a specific gravity greater than water, it will sink, although resistance to sinking

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will be offered by the viscosity of the water. The greater the surface compared to the volume the
greater will be the friction between water and body. Because of this relation, particles of very
small size, even though composed of a substance having a specific gravity greater than 1, may
not sink at all. 
Few aquatic organisms, irrespective of size, are spherical in form. Any departure from the
spherical form results in relatively increased body surface. Relative increase of surface is
accomplished in so many different ways by :
1.General body form. Main portion of body may present:
a.Various degrees of attenuation.
b.Various degrees of compression or depression or general flattening.
c. Miscellaneous forms of asymmetry.
2. Body surface sculpturing: ridges, furrows, striae, impressed or raised patterns.
3.Extensions and modifications of antennae, tentacles, gills, legs, cerci and others.
4.Development of special peripheral processes: hairs, setae, spines, bristles, filaments, radial
axes, tubercles, cilia, pseudopodia, crests.
5.Formation of colonies: linear, dendritic, radial, lamellate, irregular.
Combinations of several of these structural features in the same organism may occur,
sometimes with remarkable flotation results.
(b) In the same species
In certain plankton organisms, a striking seasonal change of body form of a very definite sort
occurs. This change of body form is a response to changes in the viscosity of the water due to
seasonal changes in temperatures.
(c) Accessory provisions of flotation
Cases and coverings of various kinds composed, of foreign materials are constructed by some
aquatic organisms. While such cases often serve several other purposes. In certain cases that
they either increase the tendency of the whole organism to be in suspension or completely
support it at the surface. Instances of this sort are not uncommon among the aquatic insects
(certain caddis-fly larvae, certain aquatic caterpillars, and others.
(d) Hydrofuge structures
Hydrofuge structures, such as hydrofuge pubescences, hydrofuge caudal filaments, and
hydrofuge smooth surfaces, often play an important and sometimes a vital part in the flotation of
organisms. Once at the surface they may provide (1) for the proper orientation of the body into
the breathing position and (2) for the ability to remain at the surface in this position without much
effort during the breathing period. Examples are not uncommon among aquatic insects. Certain
hydrofuge structures are related directly or indirectly to respiration in some air-breathing aquatic
insects.
(e) Precision in flotation adjustment
Among the plankton, flotation at the proper level is sometimes a matter of precise adjustment. A
very small discrepancy in adjustment may result in the following chamges.(1) too great
buoyancy, causing the organism to rise to an unfavorable level or even to rise to the surface.
Owing to various hazards such as excess light, entanglement with the surface film, and
evaporation; or (2) sinking to an unfavourable depth at which such features as reduction of
effective light, critical reduction or absence of oxygen, and absence of proper nutritive materials
may result seriously. Such adjustment is thus of vital importance to certain nonmotile plankters.

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This adjustment follows the seasonal changes of viscosity and density of water in such a way
that organisms may continue to thrive is one of the marvels of aquatic life.
Water fleas swim intermittently but unceasingly, and certain microorganisms occupy proper
levels by constant vibrations of flagella or cilia.
Body form adjustments
a) Streamline form
When a body is either in motion through quiet water or stationary in moving water, the water
imposes some resistance over the body. Hence, the organism has to overcome this force to
maintain its stationary position.
A body must occupy space by complete displacement of the water and that when either the
body or the water moves; the body continues the process of displacement. Also, under these
conditions, not only must the great weight (density) of the water be overcome in pushing it
aside, but the internal friction (viscosity) of the water plus the friction of the water against the
surfaces of the object must also be overcome. Thus, energy is expended by the object in
overcoming these resistances, and, as will be shown later, the amount of this energy depends
upon the form of the body when other conditions are the same.
Viscosity and density of water vary with certain conditions, particularly temperature; the
resistance met by a moving organism or by an organism maintaining its position in moving water
is considerable under all circumstances. Organisms vary greatly in their ability to overcome this
resistance, owing to inherent differences, especially the form of the body. Obviously, certain
forms of body are more effective for locomotion in water than others.
Among other things, it was found that the body consists of two principal parts: (1) the entrance
or fore body, that part from the tip of the snout to the maximum transverse section; and (2) the
run, or after body, that portion from the maximum transverse section to the tip of the caudal fin.
It was also found that, in all specimens, the average position of greatest transverse section
occurred at a distance of about 36 per cent.
The replacement of water following maximum displacement had something to do with the
function of the after body.
Principle of streamline form
A body with streamline form moving through standing water and the same body maintaining a
fixed position in running water present the same essential conditions.
The position of maximum transverse section represents the maximum displacement of water. it
likewise determines not only the maximum energy expended in displacement but also the
termination of virtually all energy expended in displacement but also the termination of virtually
all energy expenditure so far as body surface is concerned.
(b) Other forms of adjustment 
Some animals living in the strongest currents possess spines on the exposed surface
(Blepharoceridae). These projecting structures would be detrimental by increasing the
resistance. These roughnesses actually help to decrease resistance. In bodies such as spheres
and cylinders, the nature of the resistance may change markedly with relatively small changes
in the conditions involved; at certain velocities, the resistance of a sphere may be reduced by
roughening its surface. In some of the species, spines will be developed as a means of
diminishing the resistance to the fierce currents in which they live. The spines on such a body

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would increase the resistance at some water velocities but would decrease the resistance at
certain higher rates of flow.

5.3.1. Relations of organisms to movement of water, surface film relations 

Relations of organisms to movements of water
Movements of water, in the various forms, affect aquatic organisms in many ways, directly or
indirectly, and often play very important roles in aquatic environments.
a) Effects upon sessile animals
Different growth forms in the same animal are the result of presence or absence of water
currents or movements especially in fresh water sponges.
The unbranched colonies are the result of unfavorable conditions, but this can scarcely be
credited when both the branched and the unbranched forms occur side by side in the same
water and on similar supports. Bryozoa have also been supposed to develop different growth
forms in standing and in moving water.
b) Effects upon motile animals
Many motile animals show a definite orientation response to current; i.e., they exhibit either
positive or negative rheotropism. Orientation reaction may be accompanied by locomotor
activities, so that certain animals will not only head upstream but will swim, either maintaining
their orginal position or making progress against the current. Sometimes the response to current
depends upon some important event in the life history such as sexual maturity.
Atlantic smelt, established in the upper waters of the Great Lakes, which while essentially a lake
inhabiting fish, becomes positively responsive to current at the onset of spawning season and
exhibits spawning “runs” at night into certain adjacent inland waters flowing into the Great
Lakes. 
Water in motion imposes pressure against certain surfaces of the animal, and it has been held
that equality or inequality of current pressure on different parts of the body affords the stimulus
to orientation of some aquatic animals, which, if true, furnishes an instance of the direct effect of
current.
Certain fishes are supposed to orient in response to visual impressions as they float
downstream (Clausen et al., 1931) but still other fishes have been thought to orient in response
to the rubbing of parts of the body on the bottom as the current make them to floats
downstream. The visual theory seems ineffective in those instances of runs at night (smelt) or in
very turbid waters.
Certain phenomena such as morphological or physiological, may either be caused by, or
correlated with, movements of water and their different velocities. A general correlation exists
between the rate of flow and the shape of mussels in eastern Bavaria.
Current demand
Certain aquatic organisms exist permanently only in the presence of appropriate movements of
water, and it is now known that current is demanded by some of them. For example, black-fly
larvae (Simulium) of all species (one possible exception in Asia) inhabit only in rapidly running
water. Wu (1931) has shown, among other things, that these larvae possess an inherent
demand for current and that their universal absence from standing waters is due directly to the
absence of the necessary current.
Secondarily, current is also related to Simulim larvae in such matters as proper food delivery
and respiration.

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Resistance to water movement

In general, animals which meet this problem successfully do so by means of one or more of the
following features: (1) body form which offers least resistance, such as the streamline or the
hemistreamline form; (2) unusually well developed burrowing or clinging habit; and (3) special
forms of attachment to fixed, supporting objects.
A few may maintain their position because of unusually effective attachment devices (powerful
adhesive suckers of the larvae of Blepharoceridae.
Provisions for clinging and attachment
Among the numerous, special provisions for increased efficiency in maintaining position in the
face of strong water movement are the following:
1. Strong, recurved tarsal claws.
2. Exceedingly flat ventral surface.
3. Strongly depressed body.
4. Lateral margins of head and thorax produced in the form of flat margins for increased contact
with the supporting object.
5. Legs, when of large size, flattened horizontally and applied by their sides as well as by the
tarsi to the supporting object.
6. Special flattening of gills or the modification of the entire gill series to form a ventral
attachment disk.
7. Special sucking disks. Examples: blepharocerid larvae; leeches; nymphs of May fly,
Ephemerella doddsi.
8. Ventral adhesive pads, often bearing recurved spines. Examples: certain stream inhabiting
aquatic Hemiptera and May fly nymphs.
9. Terminal attachment disks. Example: Simulium larvae posterior disk a combination of a row
of hooks with a gelatinous secretion originating from the mouth.
10. Threads which anchor the animal directly to the support. Example: thread used by Simulium
larvae when shifting position.
11. Threads which anchor case or shelter of animal. Example: certain caddis fly larvae.
12. Shelters, tubes, or cases which protect against the wash of currents and waves. Examples:
sand constructed case of caddis fly, Molanna; cases of certain midge larvae; egg capsules of
leeches; tubes of tubificid worms.
13. Adhesive secretions. Example: common hydra.
Provisions for burrowing
Burrowing is often accomplished by animals having no special structural provision for that
purpose. In such instances, they are merely capable of forcing their way into bottom materials,
aided by such features as (1) more or less pointed anterior end; (2) body movements of a
penetrating sort; (3) setae; (4) longitudinal contraction and extension of a portion of the body; (5)
extensile and protrusible body tubercles; and (6) strongly muscular body walls accompanied by
freely moving, soft, internal organs and fluids. By such means as these, some of the softest
bodied aquatic animals (Oligochaeta and others) penetrate the hard packed sand of barren,
exposed shoals, thus maintaining their position in the presence of the strongest wave action.
Other animals have developed special structural features for effecting partial or almost entire
penetration of bottom, such as (1) the flattened, shovel like, anteriorly directed front legs, the
posteriorly directed hind legs deppressed to the body and adapted for pushing, and the pointed

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sloping head of the nymphs of the May flies Hexagenia and Pentagenia, and the dragon flies
Gomphus; (2) the long, upturned, mandibular tusks of burrowing May fly nymphs; (3) the
muscular foot of clams and snails; (4) the long, spraddling, spider like legs of certain dragon fly
nymphs (Macromia), so oriented on the body that they rest full length upon the sand and,
wriggling movements, work the sand entirely over them thus gaining a certain anchorage; and
(5) the strikingly flattened, shoal inhabiting dragon fly nymphs (Hagenius), which weight
themselves down by working sand on top of the thin abdominal margins.
Burrowing by some species may be a direct response to excess light, but the end result of
maintaining position remains the same.
Habits facilitating resistance to water movement
Animals, which, lacking special structural developments, manage to maintain position in current
or wave-swept areas by reactions. These are exemplified by the habitual seeking of (1) the
protected sides of and the interstices between rocks; (2) fissures in bottoms and bottom
materials; and (3) the more protected parts of rooted plants.
Influence on construction activities
Construction processes of certain animals can be properly performed only in the presence of
water movement. A striking instance is that of the net building caddis fly larvae which can
produce their nets only in moving water; in calm water, the attempt results only in a shapeless
mass of threads.
Distribution of organisms
Since moving water is an effective transporting agent, movements of water play a very active
part in the distribution of many aquatic organisms. Pieces of aquatic plants bearing various
eggs, larvae, pupae, and even adults of insects, hydra, Bryozoa, Mollusca, and many others
break from their attachments and drift with the water.
Molar agents
Sand, fine gravel, rocks of various sizes, and sometimes even boulders, carried or rolled by the
water, become a veritable wearing, grinding, fragmenting machine which constitutes one of the
serious menaces to the whole biota of those situations.
Indirect effects of water movement
Water movement is concerned with the life of aquatic organisms in a number of ways, the
following being among the most important:
1.Constant shifting of bottom materials on shoals and other shallow waters may prevent the
rooting and, therefore, the occupancy of these areas by higher aquatic plants.
2.Erosion or transportation of materials may completely alter the environment, converting it into
some very different type for which the organisms are not suitable. The example of the cutting off
of a sand-spit beach pool from the body of a lake.
3.Circulation, and in some instances the return to circulation, of essential nutritive substances in
the water, both dissolved and suspended.
4. Production and maintenance of turbidity thus affecting the light penetration and certain other
relations.
5.Delivery of food to sessile or sedentary animals, particularly when the food is in the nature of
suspended, living organisms (plankton) and suspended, finely divided, nonliving materials.
6.Respiratory relations, such as (a) renewal of properly oxygenated water to respiratory
surfaces and (b) renewal of dissolved oxygen supply from the air by the surface agitations

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incident to water movement.

7.Temporary exposure to air, as in seiches which imitate the ebb and flow of a tide and which, if
of sufficient magnitude, may expose for a time a whole set of shallow water organisms to
evaporation and other serious hazards.
Surface film relations
The surface film serves as a mechanical support for organisms and miscellaneous particulate
materials. Both surfaces of the film may function in this way. The term neuston, originally
applied to minute organisms, is now commonly extended to include all organisms associated
with the surface film. Those related to the upper surface of the film comprise the supraneuston;
those related to the lower surface, the infraneuston.
The larger animals commonly associated with the supraneuston are: (1) water striders
(Gerridae); (2) broad-shouldered water striders (Veliidae); (3) water measurers (Hydrometridae);
(4) hebrids (Hebridae); (5) mesoveliids (Mesoveliidae); (6) whirling beetles (Gyrinidae); (7)
springtails (Collembola); and (8) certain spiders.

5.3.2. Relations of organisms to Temperature relations, Light relations

Temperature relations
With the exception of the aquatic birds and mammals, all aquatic animals are cold blooded
(poikilothermous), i.e., their internal temperatures follow, usually within close limits, the
temperatures of the surrounding medium. It must be understood, however, (1) that exceptions in
the form of unusual deviations from surrounding temperatures may occur, as, for example, the
claim that certain fishes may have an internal temperature of as much as 10°C. higher than that
of the surrounding water; and (2) that the degree of agreement between body temperature and
external temperature may differ with the temperature level of the latter.
Some aquatic animals live in surroundings the temperature of which is below freezing (glacier
worms and others), but it has usually been supposed that the freezing point of their body fluids
is depressed by substances in solution. Even if this is true, there remains to be explained the
fact that under those very low temperatures they are not only active but grow, develop, and
reproduce.
Influence on metabolism
Within the ordinary temperature limits for a given cold blooded animal, decreasing temperatures
diminish metabolism, and vice versa, a relation which is opposite that for warm blooded
animals. This means that metabolic rate is, to a large extent, governed by the external
temperatures. It also means that the falling temperatures of increasing depths in water or of
increasingly northern latitudes inflict lower rates of metabolism. A general rule for this change in
metabolism in cold blooded aquatic animals can be stated as follows: a rise of 1°C increases
the rate of metabolism about 10 per cent. This means that the rate of oxygen consumption and
carbon dioxide output doubles with a temperature increase of 10°C.
Influence on development and other biological processes
Rising temperature increases the rate of (1) development of animals, (2) respiratory
movements, (3) heart beat and circulatory rhythms, (4) enzyme action, and (5) other
physiological process, although the operative limits in each process may differ. A cold blooded
aquatic animal may be expected to complete its life cycle more slowly and to produce fewer
generations per unit of time in the northern than in the southern part of its range; like wise, the
normal individual life span may be longer. Onset of hibernation, breeding season, changes in

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reproductive activity, germination of asexual reproductive bodies, and a host of other biological
activities are profoundly influenced by surrounding temperatures.
Temperature toleration
Each organism has a maximum and a minimum environmental temperature between which life
is possible but beyond which conditions are lethal. Even for individual species, these
temperature limits are not absolutely fixed, since they may vary with different individuals, with
the different sexes, with different life history stages, with different physiological states, and in
different parts of the geographic range. In spite of this variation, it is possible roughly to divide
animals into two groups: (1) those which are restricted to a narrow range of temperature change
(stenothermic animals) and (2) those which tolerate a wide range of temperature change
(euthermic animals). Also, there are integrades between these two groups. It is a well known
fact that acclimatization can shift temperature restrictions as well as those of other
environmental factors. Somewhere between the maximum and minimum limits, an optimum
region occurs, the position and extent of which vary with different animals. It is sometimes
stated that the optimum is usually closer to the maximum than to the minimum, but in some
instances the reverse condition prevails. Acclimatization may also affect the position of the
optimum.
In temperate lakes of the first and second orders, only the non-migrating, profundal bottom
organisms live under approximately a fairly even temperature throughout the year. On the
contrary, those surface water forms which remain active throughout the year must endure the
complete range of temperature. Those not active in all seasons have developed various forms
of hibernation, and aestivation, as a means of passing over the more rigorous conditions. Many
aquatic animals remain active thorough wide ranges of temperatures, the active period ending
only just before the extremes are reached.
Effects of extremes of temperature
The specific effect of extremely low temperature is usually considered as being mainly
mechanical, while that of extremely high temperature is principally chemical, affecting the
protoplasm. The chemical effect of excessively high temperature is more severe than the
mechanical effect of correspondingly low temperatures. It is true that even in the temperate
latitudes, certain aquatic animals (mosquito larvae and others) may be frozen into surface ice
and recover on release. This phenomenon seems to be more common in the arctic and
subarctic regions.
The occasional rise of surface water temperature to unusual heights (although only a few
degrees above the usual summer maximum) in protected bays in times of clear, hot weather
and dead calm water promptly leads to a dying off of surface plankton and certain other shallow
water organisms.
Recognition of temperature differences
Some aquatic animals have a well developed recognition of changing temperature and may
respond with considerable precision. Under experimental conditions, certain fresh water animals
have been found to recognize temperature differences of 0.2°C and react to them.
In thermally stratified lakes, it is very difficult to determine the presence or absence of a limiting
effect to downward distribution by the steep thermal gradient in the thermocline, since other
varying conditions are simultaneously present, such as light, chemical stratification, and
viscosity changes.

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Light relations
The various relations of sunlight to aquatic organisms may be classified into two sets: (1) direct
influences upon the organisms as a whole and (2) photosynthetic relations.
Direct influences
Lethal effects
Many aquatic organisms are sensitive to the higher intensities of sunlight and, in fact, must
avoid them by occupying deeper levels in the water.
Plankton organisms occur in surface waters, exposed to the maximum light intensity. Many are
phytoplankton and have photosynthetic relations, but it is a well established fact that in most
natural waters the maximum populations of plankton occur at some lower depth, one of the
important reasons being the more favorable light effects.
Both sunlight and ultraviolet light seemed to devitalize the diatoms (excepting Synedra) but
stimulated the Chlorophyceae and the Myxophyceae, the ultra violet light providing a slightly
greater stimulus. Since the ultraviolet light is quickly absorbed in the surface waters, its effects
are very restricted.
Many aquatic organisms, especially bottom inhabiting forms, live in conditions of almost if not
complete darkness and quickly succumb in direct sunlight. Light is often a powerful factor,
sometimes the determining one, in the distribution of organisms in aquatic environments.
Behavior and orientation
Responses of aquatic organisms are often due to, or conditioned by, light. One of the most
striking results of the alternation of day and night is the migration of certain plankton organisms
from deep water to the surface at night and their return to the depths near dawn.
Light is the principal motivating influence of this migration. For some organisms, day is the
period of general activity, night the period of quiescence; for other forms, the reverse is true.
In many aquatic animals, the light responses differ markedly with physiological state, age, life
history stage, season and other conditions.
Other influences
Direct influences of light effects upon pigments and pigment production, upon growth, upon
development, and in fact, upon many of the conditions involved in the general success of
organisms.
Photosynthesis
One of the most profound influences of sunlight (and of moonlight to a limited extent) in water is
its intimate role in the photosynthetic processes of all chlorophyll bearing, aquatic plants.
These plants furnish, directly or indirectly, the carbohydrate and the protein supply for the
aquatic world. They occupy that strategic position between the inorganic and the higher organic
components which makes the latter their complete dependents. The phytoplankton has been
called the green pasture of the sea, and it plays a similar role in fresh waters too.
It becomes increasingly clear that the process commonly referred to as photosynthesis is a very
complex phenomenon. Light, temperature, solutes, and carbon dioxide affect simultaneously the
photosynthesis. However, it seems certain that light of the appropriate kind and intensity is the
supreme factor.
Light requirements
The light supply has two important aspects: (1) light intensity and (2) effective wave lengths.
Intensity

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The rate of photosynthesis increases with the intensity of light and infact, if certain conditions of
temperature and carbon dioxide are met, the rate of photosynthesis is proportional to the
intensity of the incident light. However, the rates of photosynthesis differ in different plants.
1.Ultraviolet rays are of little or no consequence in photosynthesis. This has been demonstrated
experimentally with terrestrial plants. Considering the fact that ultraviolet waves are completely
absorbed in the uppermost, thin layer of the water and that various aquatic plants thrive far
below the level of disappearance of these wave lengths, the aquatic situation seems to offer
confirmation of the statement.
2.Experimental evidence appears to show that with equal intensity of incident light,
photosynthesis is affected by different wave lengths, being greatest in the red and least in the
blue violet. Certain investigators claim that the rate of photosynthesis diminishes with
decreasing wave length.
Effective light penetration
The normal existence of healthy chlorophyll bearing plants at various depth levels in water may
be taken as evidence that some of the effective light is present in sufficient intensity to enable
these plants to perform photosynthesis.
Algae have been found in certain mountain lakes below a depth of 400m and at greater depths
in the ocean, but it remains to be conclusively demonstrated that these plants are performing
photosynthesis.
It seems certain that light is a very influential factor in determining the occurrence and
distribution of chlorophyll in a lake. Therefore it may be expected that since light conditions differ
in different waters the quantity and activity of chlorophyll will be influenced correspondingly.
Nevertheless the processes of light penetration and photosynthesis in natural waters are so
complex.
The maximum rate of photosynthesis in lakes in full sunlight usually occurs somewhere below
the surface layer.
Plants inhabiting situations having moderately reduced light intensity usually have more
chlorophyll than do those living in full sunlight.
That light intensity at which oxygen production in photosynthesis and oxygen consumption by
respiration of the plants concerned are equal is known as the compensation point, and the
depth at which the compensation point occurs is called the compensation depth. For a given
body of water this depth varies with several conditions, such as season, time of day, degree of
cloudiness of sky, condition of the water, and taxonomic composition of the flora involved. As
commonly used the compensation point refers to that intensity of light which is such that the
plant’s oxygen production during the day will be sufficient to balance the oxygen consumption
during the whole 24hr period.
Photochemical nitrification
An indirect effect of sunlight is through a possible photochemical nitrification. A portion of the
nitrification which goes on in the sea is photochemically activated. Some chemical nitrification in
soil is activated by sunlight in the absence of the biological agencies.

5.4. Relations of Dissolved Oxygen, Relations of Carbondioxide

Relations of Dissolved Oxygen
Oxygen supply in air and in natural waters affords a striking contrast. Normally, air contains
oxygen to the extent of approximately 21 per cent, which is an abundant supply for the

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respiration of air breathing organisms.

One liter of water will contain only about 9 cc of oxygen when saturated with this gas, whereas a
liter of air will have 210 cc. In view of the active interplay of oxygen producing and oxygen
consuming processes in inland water sets the stage for serious limits in aquatic respiration.
Excess of oxygen
Moderate supersaturations of dissolved oxygen occur in natural waters from time to time,
usually owing to the photosynthetic activities of large masses of green plants in very calm water.
Under special and still rare circumstances, large accumulations of excess oxygen appear in the
upper part of the thermocline or in deeper strata of a lake.
Normal dissolved oxygen requirements
In the dynamics of natural waters, oxygen supplying and oxygen consuming processes are in
constant action, the limits of an adequate supply of dissolved oxygen for organisms become an
important matter.
The minimal oxygen requirement may be affected some what by other environmental features,
e.g., temperature, CO2 and certain conditions existing within the organism itself such as age or
life history stage. In general “dissolved oxygen at levels of 3ppm or lower should be regarded as
hazardous to lethal under average stream and lake conditions; and that 5ppm or more of
dissolved oxygen should be present in waters, if conditions are to be favorable for freshwater
fishes”. This statement assumes, of course, that other vital requirements are maintained within
their proper limits. It also applies primarily to warm water fishes. It has been claimed that cold
water fishes require a higher dissolved oxygen content.
The respiration of aquatic organisms depends not only on the dissolved oxygen content but also
in a significant measure upon the temperature of the surrounding water; that the oxygen
consumption is almost doubled by a rise of 10°C; that the same amount of dissolved oxygen
has about twice as great a supply value at 5 as at 15°C. 
Source of oxygen supply
1.Storage of oxygen
The hemoglobin of the blood may act as a storehouse for oxygen that such storage at times of
abundant free oxygen may furnish the supply during oxygen deficiency.
2.Internal chemical transformation
The idea of chemical transformations taking place within the animal, such as occur in the
utilization of foodstuffs in which oxygen is released and made available for recombination.
3.Catalysts facilitating oxygen absorption
Many profundal bottom animals have manganese in their tissues. This element may serve as a
catalyst, facilitating oxygen absorption at low tensions.
4.Atomic oxygen from decaying plant tissues
The decomposing plant tissues in the profundal mud, even under anaerobic conditions,
gradually liberate small amounts of an oxidizing substance could be utilized by the animals
living in such close relationship to the decaying plant debris.
Temporary anaerobiosis
Animals living in the muddy bottoms of shallow water or other similar conditions in which the
oxygen exhaustion occurs quickly and for limited times, may be forced to meet these
temporarily unusual conditions.
Certain representatives of Protozoa, nematodes, earthworms, leeches, and immature stages of

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insects, mollusks, fishes, and others exhibit this ability.

During oxygen lack a certain amount of energy may be released by the splitting of
carbohydrates into reduced substances, thus building up an “oxygen debt,” this debt being
repaid by the increased rate of oxygen consumption when the organism is returned to aerobic
conditions.
Some effects of insufficient dissolved oxygen
1.Attempts to migrate
In lakes, it is usually an upward migration into overlying, better oxygenated waters.
2.Onset of diseases
A close relation between insufficient dissolved oxygen to diseases of fishes, parasitic and
bacterial and serious epidemics in the fish Leucichthys artedi, which seems to occupy the cooler
water below the thermocline during summer.
3.Suffocation beneath ice cover
Shallow waters with bottoms containing large amounts of putrescible matter and occurring in
regions where prolonged ice cover in winter is common may, at times, almost or completely
exhaust the dissolved oxygen of the unfrozen water with resulting mortality (winter kill) among
the organisms.
4.Summer kill
Critically low dissolved oxygen and unfavorable temperatures have been suspected as causal
agents, but the case is not clear since other conditions were probably in a simultaneous state of
flux. Warm waters reduced solubility of DO and release of oxygen by warm waters (less holding
capacity).
Relations of Carbon Dioxide 
General effects on organisms
Carbon dioxide is one of the most important substances in the life of organisms.
Small quantities
Usually, the quantities in the air are very small but yet sufficient for the photosynthetic activities
of chlorophyll-bearing plants. Likewise, in natural waters, the amounts may be very small in the
upper circulating waters.
Large quantities
Large quantities of carbon dioxide usually have a detrimental effect. Ordinarily, accumulations in
unpolluted, natural waters do not reach such lethal amounts, owing to the ease with which they
are released into the air or combine chemically. Increasing amounts of free carbon dioxide in
association with other decomposition products may gradually render the hypolimnion untenable
by all organisms save the resistant anaerobic animals in the bottom. 
Such accumulations may render bottom waters acid in reaction and thus affect organisms
sensitive to acid waters. High carbon dioxide content seems to be more toxic in the presence of
low oxygen content.
Since an excess of dissolved carbon dioxide is usually accompanied by a much reduced
dissolved oxygen content and other important conditions, it has been proposed that the carbon
dioxide content of the water is probably the best single index of the suitability of water for
fishes. 
Carbon dioxide has a very definite effect upon the affinity of blood for oxygen in fishes and
certain other animals.

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Fishes may tolerate wide, but not sudden, ranges of carbon dioxide tension of the water by
“increasing the alkali reserve of their blood in high carbon dioxide tension water and by lowering
the alkali reserve of their blood in low carbon dioxide water”.
Carbon dioxide tension
While the carbon dioxide tension within natural waters and within the atmosphere constantly
tends toward equilibrium, circumstances prevailing in the water (slow diffusion, rapid production
of carbon dioxide, insufficient agitation of the water) may be such that at some depths the
carbon dioxide tension is greater than in the air.
Relations of other Dissolved Gases : Methane, Hydrogen Sulfide 
Methane
Some have claimed that it is nontoxic; others, that its effects on organisms are minor or that, at
most, it may be occasionally toxic to animals or that it may cause them to migrate from
particular situations. Possibly methane accumulations in bottom waters may have something to
do with the increasingly severe conditions which develop with the progress of stagnation
periods.
Hydrogen sulfide
Inherently, hydrogen sulfide is very poisonous. Certain marine fishes are said to be very
sensitive to this gas and to avoid water containing it.

5.5. Nitrogen, Ammonia, Dissolved solids, Other elements, Dissolved organic matter
Nitrogen
Free nitrogen has usually been supposed to be the least important of the dissolved gases when
it occurs in normal quantities. Excess nitrogen is said to cause gas disease in fishes. Unusual
amounts may produce entry of the gas into the circulatory systems of aquatic animals, causing
stoppage. 
Ammonia
Scanty information is available on the biological relations of gaseous ammonia as produced in
natural waters.
Dissolved solids
Relations of inorganic nitrogen compounds
Ammonium salts, nitrites, and nitrates furnish a supply of nitrogen which is essential in the
fundamental food relations of organisms. Ammonium salts (“ammonia nitrogen” or “free
ammonia”) constitute the first stage in mineralization of organic nitrogen. It is usually considered
that nitrates supply nitrogen in more available form, although the other two compounds,
particularly ammonium salts, are utilized to some extent.
While some plants seem to prefer nitrates, there are others which grow equally well with both
nitrites and ammonium salts. Variations in the quantities present in water are correlated with the
growth seasons of plants and with the temperatures which control, to some extent, the rate of
bacterial action. Ordinarily, nitrogen in its final oxidized form as nitrate does not occur in great
amounts in natural, uncontaminated waters. The Algae, water weeds, and nitrate reducing
bacteria are the important consumers of nitrogen content and that the nitrifying bacteria aid in
increasing the nitrate content. Nitrogen is considered to be one of the most important limiting
factors in the development of phytoplankton. It is one of the nutritive substances necessary for
the production of chlorophyll. Formation of chlorophyll ceases very quickly with nitrate
deficiency. Ammonium salts in excess are reported as poisonous to fishes if present with

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carbonates.
Relations of silicon
Since diatoms require silicon for the manufacture of their shells, and since they constitute a very
prominent and strategic group in the plankton at large, the available supply of silicon in the
water is regarded as a matter of importance. The production of diatoms is directly determined by
the silicon supply. Silica deposition by diatoms is a one way process; that silica in the form of
diatom shells is highly resistant to passage into solution in water; that diatom shells once formed
are practically permanent in many waters. Silicon removed from sea water by diatoms and other
organisms may return to solution after they die, or it may sink to the bottom. 
Development and success of the fresh water sponges depend upon an adequate supply of
silicon for the manufacture of spicules. Some permanent loss of silicon is expected, in average
situations, owing to transportation by currents, to outlets, and to burial in bottom deposits. In the
presence of such losses, a source of renewal is necessary if a body of water is to avoid silicon
decline to a critical level.
Relations of phosphorus
Since the amount of soluble phosphorus in natural, unmodified waters is small and since
phytoplankton requires an adequate supply of phosphorus, it is now generally regarded as a
limiting factor. Phytoplankton occupies the upper waters because of their light requirements die
and sinks to the bottom, carrying away a certain amount of the phosphorus. Restoration of
phosphorus to the upper waters might be brought about by inflow of waters rich in phosphate or
by the return to circulation of the phosphorus containing materials, return of which would be
facilitated by overturns or other forms of circulation.
The nitrogen phosphorus ratio
The relation of these two substances is better known for sea water than for fresh water. The
concentrations of the two substances closely parallel each other. In sea water it appears that
the ratio tends to approach a constant value, with nitrogen exceeding considerably the
phosphorus content, and that as claimed these substances occur in marine plankton in about
the same proportions. Further, the ratio in inland waters may be different not only in numerical
value but also in the range of deviation from proposed mean. This matter is still in the
pioneering stage, but is suggestive of basic limnological possibilities and deserves more
investigation.
Other elements
The significance of several other elements appears most prominently in their essential roles in
the metabolism of the various groups of aquatic plants. Calcium is required by all green plants
except some of the lower Algae; is not necessary for the fungi; and while necessary for the non-
chlorophyll flowering plants, they usually contain less calcium than do chlorophyll bearing ones.
It appears to have several physiological roles, such as (1) relation to the proper translocation of
the carbohydrates; (2) an integral component of plant tissue; (3) facilitation the availability of
other ions; and (4) an antidoting agent reducing the toxic effects of single salt solutions of
sodium, potassium and magnesium.
Magnesium is a component of chlorophyll and must be present for its proper development. It
appears to act as a carrier of phosphorus, at least in some instances. Quantities of magnesium
larger than usual in natural waters may be toxic to some aquatic organisms. Cladocerans are
wholly absent from certain lakes (Lake Tanganyika, Lake Kivu) may be owing to the excess of

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magnesium over calcium salts.

Iron must be supplied for plant growth and development. It functions in the proper production of
chlorophyll, although it does not enter into the chemical composition of chlorophyll. It acts as a
catalyzer; others, that iron is the oxygen carrying substance in certain respiratory processes.
Both the quantity and the form in which it is presented to the plant are now known to be
important, these being conditioned by the features of the environment (hydrogen ion
concentration, organic matters, and others) and the kind of plant involved. Most algae grow best
when the water has a ferric oxide content of 0.2 to 2 mg/litre, but distinct toxicity occurs when
the available iron exceeds 5mg. however, many natural waters may contain more than 5mg. of
iron without being toxic owing to the buffer action of organic compounds or of calcium salts.
Toxic oxidation products of pyrites are said to be formed in peat deposits. Two other relations of
high iron content have received attention in limnological literature: (1) reduction of nitrates to
nitrites by ferrous salts in the presence of oxygen and (2) reduction of dissolved oxygen in the
presence of iron.
Sodium, while apparently not absolutely necessary for plant growth and development, is
evidently a very desirable element. It may serve one or more of the following roles: (1) act as a
conserver of potassium, since less is absorbed when sodium is present; (2) replace potassium
to a limited extent as a plant nutritive element; (3) render soil-absorbed potassium more
available to plants; and (4) be an antidoting agent against certain toxic salts in the medium.
Potassium is a fixed requirement for plants. Its function is imperfectly known, but it appears to
be two fold; (1) a fundamental requirement in food manufacture and (2) a catalyst. Sulfur must
be provided for plant growth and development. It forms a necessary material in the composition
of protein and other constituents of the plant.
Trace elements
By trace elements is meant those chemical elements essential to the well being of animals and
plants but required only in extremely small quantities. Prominent among these trace elements
are copper, manganese, zinc, boron, lead, cobalt and iodine. 
Dissolved organic matter
Many of the minute, more or less undifferentiated organisms, such as the bacteria, certain
Algae, and certain protozoa, must depend upon the dissolved materials in their environments for
the substances necessary to growth and development. At one time, the discovery that many
microorganisms use particulate foods cast some doubt on the direct utilization of dissolved
substances by many organisms. However, it has since been clearly demonstrated that bacteria,
diatoms, most if not all other phytoplankton, and some of the Protozoa normally utilize and
depend upon both the dissolved inorganic and the dissolved organic materials in their
surroundings. It likewise seems probable that many other small organisms which lack a
digestive tract or other provisions for introducing particulate foods will be discovered to depend
upon these materials, wholly or in part.
The plankton in the ocean is entirely insufficient in amount to supply the necessary nourishment
for those animals supposed to depend upon it and (2) that an abundant supply of food is
available in the dissolved organic matter in the water. From computations of the minimal carbon
requirements of certain marine animals for a given time unit and determinations of the plankton
content of the surrounding water, he insisted that the amount of water strained by the animal to
secure the minimal amount of carbon was impossibly large. In this way, he showed that a

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certain common marine sponge would need to filter 2421 of sea water per hour (about four
thousand times its own bulk) to secure the minimal amount of carbon from the plankton.
The chlorophyll bearing phytoplankton is not dependent upon nitrogen salts or upon carbon
dioxide in water for either nitrogen or carbon but, instead can utilize atmospheric nitrogen
dissolved in the water (nitrogen fixation); that the nitrites, nitrates, and ammonium salts in the
water may remain unconsumed and that bicarbonates of calcium and magnesium can be
broken up, the half bound carbon dioxide furnishing a carbon supply for the green
phytoplankton. This reaction, they claimed, is of such magnitude that at the spring plankton
maximum, assuming that it occurs to the same extent down to a depth of 100m., the carbon so
provided would be sufficient for a phytoplankton crop of 10 tons or more per acre, wet weight.
1.Organic detritus and living organisms in water usually provide food in necessary quantity for
the aquatic animals present. Certain protozoa and possibly sponges may absorb dissolved
organic matter from the water. 
2.The rather large quantities (10 mg. per 1. or more) of dissolved organic matter in fresh water
include proteins in colloidal solution and several amino acids greatly diluted. Carbohydrates
present do not appears to be in readily assimiable form.
3.Dissolved organic matter seems to be principally waste products, some of which are very
resistant to bacterial action.
4.Very little of the organic matter produced by living Algae is given off to the water; 90 to 95 per
cent of it is stored in the organisms.
5.It is possible that higher animals absorb insignificant quantities of the dissolved substances. 
6.Experimental evidence is now available which indicates that tadpoles, mussels and probably
other animals may take up dissolved organic matter from rather concentrated solutions and are
thus enabled to thrive and grow, at least for a considerable period of time, in the absence of
particulate food.
7.Experiments of certain investigators show that absorption of dissolved organic matter by
tadpoles, mussels and starfish occurs through the intestine and not through gills or integument.
The integument and gills of aquatic animals seem to be, for the most part, impermeable to
organic substances.
In general, marine invertebrates are permeable to water, salts, and organic solutes but that
teleosts and fresh water invertebrates are very slight permeability. In Daphnia magna organic
substances in true solution were not used for food, but organic matter in colloidal form was
utilized. The fishes may absorb a slight amount of dissolved substance, but the securing of a
large proportion of their nutriment in this way, as has been postulated, appears very doubtful. 

Unit 6- Planktonic organisms

6.1. Classifications of organisms in water
Planktonic organisms
The term plankton was first proposed by an Oceanographer, Victor Hensen in 1887 to designate
that the heterogeneous assemblage of minute organism and finely divided, non-living materials
which are known to occur in the waters and to float at the will of the waves and other water
movements. 
Classifications
Plankton of the various freshwaters differs widely in quality. The following outline is an
abbreviated indication of the organism which may occur in the plankton of fresh water.
1. Plants :

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a) Algae are represented in the plankton of inland waters

b) Fungi, which occur as bacteria abundantly in the plankton, in fact it is likely that no water in
nature is free from them.
2. Animals :
a) Protozoa – These are the representative of the plankton by many genera and species 
b) Coelenterate - Mainly hydra is facultative plankter occurring at times free in open water 
c) Rotatoria is the most important groups of zooplankton
d) Gastrotricha - occur in limited numbers 
e) Bryozoa whose larvae are free swimming 
f) Arthropoda – Mainly i) Crustacea and ii) Insects 
3.Occasional plankters : 
a) Flowering plants for eg. Wolffia (Lemnaceae) occur at various depth and are recorded as
plankton organisms especially in rivers.
b) Platyhelminthes eg, Turbellaria are of less importance in freshwaters.
c) Coelentrata eg, Medusa (Craspedacusta) 
d) Insecta eg, May-fly nymphs
e) Arachnida eg, Water mites
f) Vertebrata eg, Juvenile stages of fishes
Classification and Terminology of Plankton
Some of the important groups of plankton are :
1. Classification based on Quality
A. Phytoplankton – plant plankton
a. Phytoplankton proper – chlorophyll bearing plankton
b. Saproplankton – bacteria and fungi
B. Zooplankton – animal plankton
2. On the basis of Size
A. Meroplankton – the large units of plankton, visible to the unaided eye
B. Net plankton (Mesoplankton) – plankton secured by the plankton net equipped with No. 25
silk bolting cloth (mesh 0.03 to 0.04 mm)
C. Nanoplankton (Microplankton) – very minute plankton not secured by the plankton net with
No 25 bolting cloth
a) Ultra plankton : <5 µm (<0.005 mm)
b) Nano plankton : 5-60 µm 90.005 – 0.06 mm)
(Dwarf / runts)
c) Microplankton : 60-500 µm (0.06-0.5 mm)
(Net plankton)
d) Meso plankton : 500-1000 µm (0.5-1.0 mm)
e) Macroplankton : 1.0 mm – 1.0 cm
f) Megaloplankton : > 1.0 cm
3.On the basis of local environmental distribution
A. Limnoplankton : lake plankton
B. Rheoplankton (Potamoplankton) : running water plankton
C. Heleoplankton : pond plankton
D. Haliplankton : salt water plankton
E. Hypalmyroplankton : brackish water plankton
4.Based on Origin
A. Autogenic plankton : plankton produced locally
B. Allogenic plankton : plankton introduced from other localities
5.On the basis of content
A. Euplankton / True plankton
B. Pseudoplankton (Flase plankton) : debris mingled in plankton

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6.Based on the life history / plank tonic life (Length of time)

A. Holoplankton / Permanent plankton : organisms free floating throughout their life
B. Meroplankton / Temperory plankton : organisms free floating only at certain times or stages
of life cycle
7.Based on habitat in water body
A. Hypoplankton : Benthic
B. Epiplankton : Surface
C. Bathyplankton : Aphotic zone
D. Mesoplankton : Disphotic/lighted zone
6.2. Distribution of plankton
In all natural waters irrespective of latitude, longitude and physic-chemical characters majority of
normal supporters are of plankton and differ in great many respects. Occasionally they occur in
certain thermal waters, subterranean waters, spring-fed streams, transient pools etc. The
presence of plankton in natural waters is itself an indication of the significant position which it
occupies in the aquatic environment.
General geographical distribution of plankton 
It has been claimed by some of the Oceanographers that the polar seas support abundant
plankton than do the tropical ones. Polar and tropical inland water are still little known
limnologically plankton production cannot be compared satisfactorily. Certain northern low lying
lakes in countries bordering on the Baltic sea, plankton production in the inland water of North
American is little known. 
Horizontal distribution of plankton
There is irregularity in horizontal distribution of plankton in water. One of the most common
causes of irregularity is the wind acting upon surface waters. Plankton drifts due to wind action
are always temporary and is common in many inland waters. This drifting leads to meeting
towards the shore becoming so thick that the whole water colour is altered and changes the
general appearance. The general effect of drift is to concentrate more or less upper waters
throughout one part of a water body with corresponding thinning on the opposite side.
Other important factors which are responsible for horizontal distribution of the plankton are :
a) inflowing streams
b) irregularity of shore line
c) depth of water
d) flowage areas
e) water current etc. 
Wind is one of the most common causes of irregularity in horizontal distribution of plankton.
Since wind causes waves but in addition may produce an actual drift of the upper waters. But
under certain conditions of drifting water, plankton organisms become concentrated temporarily
in the vicinity of the shore which faces into the wind at that time.
Vertical distribution of plankton
The vertical distribution of plankton is a complicated matter. In deeper region, plankton may
show little or no resemblance to that of upper waters. In lakes, a uniform vertical distribution of
plankton occurs only during Spring and Fall overturns but certain plankton are distributed in
minimal quantities from the surface to the lowermost limits of habitability in the range of
concentration that they occur.
Distribution of phytoplankton 

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The upper most waters are the home of the chlorophyll bearing plankton and perhaps the light
plays an important role indicates the distribution of phytoplankton. Considering the general
mass distribution, few stalemates are commonly taken such as :
a. The blue green and green phytoplankton (Myxophyceae and Chlorophyceae) concentration is
maximum than diatoms and this has been thought to be due to the heavier weight.
b. Maximum populations of chlorophyll bearing phytoplankton are at some level below the
surface waters.
c. The blue green algae as a group tend to concentrate towards the surface.
Distribution of zooplankton
It is difficult to generalize on the vertical distribution of zooplankton as a whole and the same is
true of the various taxonomic groups composing the zooplankton. Certain tendencies in vertical
distribution appear such as :
a. Greater occurrence of Sarcodina in lower waters
b. Preference of Dinoflagellata for upper water
c. General scattering of Ciliatei and
d. Selection of different levels by the young and adult stages of certain crustacean
The distribution behavior may be very different in different kinds of water. Under conditions of
well developed physic-chemical stratification, the levels of maximum population of the
Crustacea and Rotifera often correspond closely, although such a statement cannot be
dependent upon.
Conditions influencing vertical distribution
Among the influences which may operate in the production of various forms of vertical
distribution, the following are important a) Light b) Food c) Dissolved gases d) Temperature e)
Wind f) Gravity g) Age of individuals of a species.
1. Light is the most important factor in vertical distribution of plankton and is well established.
Its presence for various reasons tends to removal or sometime disappearance of plankton. The
annual and diurnal variation of light, qualitative and quantitative variation influence the migration
of plankton Differences is light reaction of young or adult stages of certain plankters lead to a
different vertical distribution of the life history stages in the same species. 
2. Food - The abundance and availability of zooplankton also depends to same extent by the
distribution of food. It has been wall established that the concentration of protozoa, micro
crustacea and rotifers have been correlated with the presence of settling suspended materials. 
3. Dissolved gasses and other substances - Dissolved oxygen is an important part in rendering
deeper waters partially or wholly uninhabitable for most of common zooplankters. Some
zooplankters are more sensitive than others to the accumulating results of decomposition in
underlying waters so that all which do not retreat upward at the same rate and certain
stratification may result. But nevertheless, the chemical changes progress to the lethal point for
pelagic plankton organisms eventually eliminating from underlying waters. 
4. Temperature
Direct effects : 
a) selection by motile plankters of favorable temperatures or 
b) inability of non-motile forms to exist in levels having certain temperatures.
Such effects appear to apply only to those plankters which manifest sensitively to differences in
temperature while many plankters are not influenced at all by any of the vertical temperature

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differences within a lake.

Indirect effects : changes in density and viscosity of water altering the floatation levels of those
plankters which are adjusted to floatation.
5. Wind – its effect is significant with the season. During summer, directly influence the
epilimnion and during hard blows vertical distribution of plankton is seen. During Autumn and
Spring overturns, wind disturbs plankton because of same density and viscosity throughout but
during the period of ice cove, the disturbing influence of wind is eliminated
6. Gravity – reduction of specific gravity makes certain phytoplankton such as Gloeotrichia to
congregate at surface waters. Plankton especially the pelagic crustacean are heavier than water
and sink with appreciable speed when inactive. Daphnia maintain their position in water.
7. Age of individual of a species – as a general rule, young individuals occur near surface,
adults tend to sink into deeper waters

6.3. Food for plankton organisms

Plankton contains miscellaneous assemblage of organisms which are fairly high in animal scale,
food requirement are diverse and can be considered best in connection with individual group.
Green phytoplankton: Chlorophyll bearing phytoplankton utilise organic and inorganic materials
dissolved in water. Certain chlorophyll bearing Protozoa utilize certain amount of particulate
materials.
Non green phytoplankton: They do not possess chlorophyll such as bacteria depend upon
dissolved materials.
Protozoa: The protozoan plankters feed upon minute algae and bacteria. Utilisation of dissolved
substances has been demonstrated in certain protozoa.
Metazoan – Neumann (1929) considered the relation of zooplankton as seston feeders into the
4 types viz.
1. grasping type : such as Rhizopoda which touch and secure seston with pseudopodia
2. filtration type : filtering of seston from water as animal moves 
3. sedimentation type : capture of seston by means of induced water currents and
4. predatory type : capture of other organisms 
Cladocera: They obtain food by active filtration of water and by predation. Cladocerans filter
their food from water mainly particulate matter such as inorganic debris, organic debris and
living organisms. Experiments showed that Daphnia and Bosmina took ordinary pond debris,
algae, carmine and finely divided humus indiscriminately filling the digestive tract with mixture.
Copepods: Thorough examination of the digestive tracts of certain copepods have revealed
that 
a) Cyclops had fragments of exoskeleton of Entomostraca, jaws of rotifers
b) Diaptomus and 
c) Nauplii had eaten finely granular mass for minute algae.
Seasonal Changes of Body Form in Plankton organisms
Morphological character of body form changes 
The seasonal change of body form differ greatly in different plankton and the changes from
winter to summer forms by way of increase in body surface compared to body volume.
For eg, 
a) the flagellate shows a tendency in summer to have a longer stalks
b) the Ceratium hirundinella develops in summer and produce a longer, narrower body from

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c) Cladoceran body in summer is higher than long with a longer back and longer posterior spine
and is reverse in winter season
d) Rotifers show changes in body size, changes in number and development of anterior spines
in pond condition which is less frequently in lakes.
General biological features
The following statements are based largely upon the summary by Wesenberg Lund (1926)
1. Seasonal changes of form is best developed among perennial species during summer and
disappear during winter 
2. Seasonal changes of form may take a different aspect in different lakes due to local
variations 
3. Transition ie, transformational stages occur almost abruptly over a period of 2 or 3 weeks in
autumn but is gradual from summer to winter periods 
4. The body form may not change in different season if it is having same body shape in
summer 
5. Body form among cladocerans and rotifers is restricted in case of females
Primary productivity
The primary production in the aquatic ecosystem starts with the synthesis of organic
compounds from the inorganic constituents of water by the activity of plants / phytoplankton in
the presence of sunlight. The inorganic constituents which form the raw material for this
synthesis are water, carbon dioxide, nitrate ions, phosphate ions and various other chemical
substances. The products are mainly carbohydrates and proteins and fats in very small
quantities. Organic production by plants is the first step in tapping energy by living beings from
non-living natural resources and hence called primary productivity.
The method of estimating primary productivity by dark and light bottle method was introduced by
Garder and Gran (1930). In this method, the water samples are incubated for a certain period in
light and dark bottles which are then suspended at the same depths where from the sample are
taken. In light bottles, oxygen is released as a result of photosynthesis and a part of oxygen is
used for community respiration. In the dark bottles, only oxygen consumption takes place as a
result of respiration. The amount of oxygen liberated by phytoplankton during photosynthesis is
considered as a measure of primary production.
The total quantity of organic matter synthesized by a unit measure of plants in a unit time is
termed as gross primary productivity. Some of this material is broken down by plants
themselves for their respiration, excretion and death etc. The reminder which becomes the plant
tissue is called net primary productivity.
Factors influencing primary productivity
The rates of photosynthetic primary production by phytoplankton vary greatly in different waters
and at different times. The variation in photosynthetic rates suggests that there are factors that
differ from place to place and from time to time, which determine the evident differences in
photosynthetic activity. The major influencing factors are :
1. Light intensity
The radiant energy that reaches the surface reacts with the dissolved and particulate materials
present in the water and this reaction brings about absorption and scattering. Illumination of
surface layers varies with place, time, light intensity, water transparency, diurnally, seasonally
and attitudinally and also with cloud condition and atmospheric absorption. Depending on the

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conditions, about 50% of the incident light is reflected back; about 80% of the total radiation
entering the surface is absorbed within the upper 10 m. and only about 0.1 to 0.2% is converted
into photosynthetic production.
2. Temperature level
Very high temperatures inhibit photosynthesis since they damage the enzymes and cell
structure as in photoinhibition. Relatively hot and light surface layer is more vulnerable to
turbulent mixing due to which the algae can be carries down below the photic zone. At critical
depth, the total primary production in the water column above equals the total loss by respiration
in the same column.
3. Nutrient supply
The fundamental importance of nutrients is that the rate at which they are supplied may
determine the rate of primary production. The potential limitations of producer activity by
nutrients show that after addition of nutrients, net production increases. The productivity of the
system is nutrient limited regardless of the changes in species composition that often result from
the enrichment.
4. Grazing rate
The biomass of zooplankton generally coincides with minima of phytoplankton density on
account of grazing. In some areas of the water bodies, as much as 99.5% of the net primary
production may be grazed. The plankton upon death, would liberate phosphorous and nitrogen
rapidly in the water making it available to phytoplankton growth.

Unit 7- Aquatic plants

7.1 Aquatic plants- Character, classification, zonation, seasonal relations
Character of larger aquatic plants 
Larger aquatic plants constitute a heterogeneous group composed of a few Bryophytes and
Pteridophytes and many of the families of Spermatophytes. They are restricted in distribution to
the general vicinity of the shores and to the shallow water areas. The larger aquatic plants of
United States are cosmopolitan within the continent although they are highly specific in habitat
requirements.
In general, the aquatic plants can be grouped in to 3 assemblages
1. Emergent – those rooted at the bottom and projecting out of the water for part of their length
eg, common species of bulrush, Scirpus
2. Floating - these which wholly or in part float in the surface of water and often do not project
above it eg. Duck weed Lemna and
3. Submerged - those which are continuously submerged eg, Vallisneria.
More or less elaborate grouping or classifications of larger aquatic plants have been proposed
from time to time. Hence, the following classification taken with some modification from Akber’s
work on Water plants in 1920, has many point of usefulness. 
Biological classification of the larger aquatic plants 
A.Plants roots in the bottom 
1) Terrestrial plants capable of living at least temporarily as submerged water plants without any
marked adaptation of leaves to aquatic life eg: Achillea ptarmica, Nepeta hederacea 
2) Plants sometimes terrestrial, sometimes with submerged leaves but different from aerial type
and associated with flowering stage
3) Plants which produce 3 types of leaf (a) submerged (b) floating and (c) aerial

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4) Plants which in certain instances may occur as land forms but normally submerged and
characterised by a creeping axis bearing long, branching leafy shoots with no floating laves 
a. Leafy aerial shoots produced at flowering period eg, Myriophyllum 
b. Inflorescence raised out of water but not aerial foliage leaves eg. Potamogeton, Myriophyllum
c. Inflorescence submerged but organs raised to the surface eg. Anacharis
d. Inflorescence entirely submerged eg. Najas
5) Plants which may occur as land forms but commonly submerged characterized by an axis
forms with linear leaves arise
6) Plants which are entirely submerged having vegetative either root or shoot naturally attached
to the substratum 
B. a) Plants which are not rooted to the bottom but live unattached in the water 
1) Plants with floating leaves, flowers raised into the air and roots not penetrating the bottom eg.
Spirodella, Lemna 
2) Rootless eg. Wolffia 
b) Plants entirely or practically submerged; Rooted but not penetrating bottom eg. Lemna and
Rootless eg. Utricularia, Ceratophyllum 
Zonation 
It is the most noticeable features of the larger aquatic plants is the distinct tendency in more or
less parallel zones along the margins of lakes ponds and similar body and water.
Smaller lakes or the protected area of the water body and marginal regions of larger lakes the
zonation may exhibit great regularity. A lake margin possessing this typical zonation shows the
following sequences :
1) Zone of emergent hydrophytes – Here plants are rooted to the bottom, submerged at the
basal portions and elevated into the air at the tops. It is a shoreward zone extending from near
the edge of water lakeward up to a depth of 2 m. Most commonly found plants include Scirpus
(bulrush), Typha (cat tail), Sagittaria (arrow heads) etc.
2) Zone of floating hydrophytes – This zone occurs beyond (lakeward) the emergent zone
comprising plants which are rooted to the bottom but the foliage float on the surface. The depths
occupied somewhat but usually about 10 cm to 2.5 m. The characteristic plants are water lilies
Nelumbo, Nymphaea, Nuphar, pond weed Potamogeton.
3) Zone of submerged hydrophytes – Typically, this zone occupies deeper water beyond the
zone of floating plants extending downward to a depth not exceeding 6 m. The plants of certain
species of pond weed (Potamogeton), water milfoil (Myriophyllum), Vallisnaria, Najas and others
are commonly found in this zone.
Seasonal relations
In regions of well defined winter season accompanied by the development of ice water and
plants of emergent and floating types die down and disintegrate with the onset of winter. This
disintegration occurs among the wholly submerged plants and still uncertain and evidences
have shown that atleast few of them were in active condition throughout the year even under the
ice.

7.2 Quantities produced, chemical composition, distribution in different waters,

limnological role
Quantities produced
Production of aquatic plants varies greatly with the nature of the water. It has been studied in

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lake Mendota, the Vallisneria constituted about 1/3rd of the total quantity; while in Green lake it
composed < 10% where Chara comprised roughly one-half of the total quantity whereas in lake
Mendota it amounted < 5%. Therefore it has been predicted that the production of larger plants
differ widely both qualitatively and quantitatively.
Chemical composition
During the growth period of plants, temporarily certain essential substances from water and
bottom deposits the chemical composition of plants which may give some information on
amount of substances used. Plants make demand upon the supply of essential materials in the
water. The amount of different substances removed or returned to the water have been studies.
The most important ones are Ash, crude protein, ether-extract, fiber and carbohydrates have
been determined.
Distribution in different waters
It is well known fact that the chemistry of water influence in determining general distribution. The
qualitative composition of aquatic flora differs in different types of water bodies in inland areas.
The aquatic flora has been divided into 3 major groups namely
1. soft-water flora
2. hard-water flora and
3. alkali or sulfate-water flora
Limnological role
It is necessary to consider the functions which the larger aquatic plants play in aquatic complex.
The relations, direct and indirect are numerous but the following ones are probably the most
important.
a. Utilization of non-living matter
The utilization of mineral salts and CO2 in the building up of green plant tissue needs more
emphasis here. It has been reported that the roots of larger aquatic plants serve primarily as
provisions for holdfast having very little physiological function in absorbing nitrates from bottom
and the absorption of nutrient materials is performed mostly by the body of the securing
substances from water only. Some of the plants roots are very much reduced and the function
of absorption is insignificant. However, the function of anchorage has an outstanding role in
these plants and like wise the reduction of absorption by the roots compared with land plants is
also established.
b. Food for animals
The role of the phytoplankton in basic food supply has been known but different opinion existed
as for as larger aquatic plants concerned as food materials for aquatic plants. Recent studies
have shown that large quantities of these plants are often consumed by great variety of animals.
Berg (1950) studied that about 17 species of Potamogeton have been consumed by 2 dozen
different species of insects. Fragments of larger aquatic plants occur in stomachs of fishes.
Analysis of stomach contents have been reported in certain fishes the plant materials
composing up to 50% of the total food content and hence it is mentioned that some fishes are
largely plant feeders. Aquatic birds (ducks, geese), mammals (musk rat, deer, moose and
beaver) secure food from aquatic plants. Nelsom et al. (1930) found that some of the plants like
Anacharis, Myriophyllum and Vallisneria have high protein and carbohydrate content including
several vitamins.

Unit 8- Nekton

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8.1 Nekton- Composition, Distribution, Movements 

Nekton 
The term nekton has been coined by Ernst Heckal (1890). Nekton is derived from Greek word
means swimming. The term nekton is used to designate those organisms which swim freely in
water and possess locomotion enabling them to have independent drifting movement along the
water flowage system. In limnetic regions of inland waters, the nekton is composed almost
entirely of fishes.
The limnetic nekton may inhabit the whole of open water of a lake and down to its greatest
depth. In contrast to the limnetic region, the littoral area is the zone of greater nekton population.
In addition to the fishes (young and mature), numerous free swimming invertebrates occur
mainly insects. Vegetative zone particularly the pondweed zone, contain largest nekton
population. Major groups of vertebrates including fishes, Amphibia, Retillia, Aves and
Mammalian are more or less representative of water and some time free swimming in nature.
Composition: Nekton comprises three phyla, viz, chordates, molluscs and arthropods, but
molluscs composition in fresh water is very less in the case of nekton.
Among Arthropods the dominant groups comprising Insects and crustaceans are involved.
However, the phylum, Chordata, is the largest group in Nektonic community, mainly fishes,
Amphibians, Reptiles, Birds and mammals.
Distribution
Distribution depends on the following factors
- seasonal changes
- Physiological changes (competition for space).
- Food availability and distribution
- Life cycle - young ones of white fishes goes to the surface for feeding whereas adults inhabit
the column water.
- Formation of local aggregations.
- Seasonal migration
- Diurnal movement
- Reproductive cycle
- Presence or absence of shelter
- Oxygen distribution
- Temperature conditions. 
It has been observed that certain fishes select deeper, cooler waters, intermediate regions and
some select upper strata. Such a distribution depends upon the season, physiological state of
the fish and the stage in life history. According to Shelford (1913) in Great lakes, the white fish
exhibit horizontal depth stratification and are arranged one above the other in their distribution.
Koelz (1929) claimed that this fish in Great lakes have shown vertical distribution pattern. In
some lakes the species that regularly inhabit shallow water may be driven by competition on the
shoals or by absence of shoals found in the deeper waters eg, lake Ontario. In lake Nipigon the
species inhabiting deeper waters are known to occur in shallow waters are Clupeaformis,
alpenae, zenithicus etc; but Johannae, Nigripinnis and Kiyi found in deeper waters.
The food of great lakes white fishes seams to be secured from the plankton and the immature
fish come to uppermost waters giving them a vertical distribution from that of mature stages.
Fishes like lake trout (Cristivomer namaycush) and turbot (Lota lota) occur in deeper waters.

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Like plankton, the nekton also have uniform distribution pattern. They have a complex
background of distribution with regard to seasonal migrations, diurnal movements reproductive
cycle, movement of water, temperature conditions etc.

Unit 9- Benthos
9.1 Classification of benthic regions, beach zones, periphyton,distribution
Benthos
The term benthos includes all bottom dwelling organisms, comprising the great assemblages of
plants and animals. Benthos includes the organisms of the bottom from uppermost water
bearing portion of a body of water right up to the greatest depths.
Classification of benthic regions
The benthic habitat is not uniform and it can be classified into different zones, they are.
1. Lake zone:
There are usually 3 zones and in exceptionally in deep lakes there are 4 zones, they are Littoral,
Sublittoral, Profundal and Abyssal zones.
i) Littoral Zone : Extends from water’s edge to the lakeward limit of rooted vegetation.
ii) Sublittoral Zone : From lakeward limit of vegetation down to about the level of the upper limit
of the hypolimnion.
iii) Profundal zone : The entire lake floor that bounds the hypolimnion 
iv) Abyssal : Present only in lakes of depths greater than 600 m.
1.Beach zones
Benthic region was formerly regarded as water’s edge extending to the deepest region. The
biota is truly aquatic, living in water at beach and has direct continuity with the main body of the
lake or stream and the organisms involved are benthos and the environment are considered as
benthic region. The exposed sandy beaches maintain 3 parallel environments, they are :
a. The inner beach
b. The middle beach
c. The outer beach
The inner beach extends from water’s edge, during periods of calm water up to the slope to the
place where the surface of sand ceases to be with water and shows the first signs of drying.
This zone is narrow and exposed to the slight wash of gentle wave of calm weather.
The middle beach occupies the space just beyond the inner beach to the waves during the
rough weather. The outer beach extends from middle beach to the outer limits of beach proper.
It is washed by waves during storms or during times of highest water levels.
2.Sandy Beach (Psammolittoral)
This is divided into three categories, they are as follows :
i. Hydropsammon : Submerged sandy bottom
ii. Hygropsammon : Zone immediately landward from water, edge – always saturatd with water
roughly corresponds to the inner beach.
iii. Eupsammon : Zone above hygropsammon – corresponds to middle beach.
Benthic Communities of Inland waters
The benthos of inland waters are vast assemblage of flora and fauna. However, it must be
remarked that, “no vertebrates are represented as true Benthos”. The benthic organisms may
be classified as follows :
i) Based on size

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a) Macrobenthos (more than 0-5 mm) eg. Crab, insects and their larvae, other crustaceans,
molluscs such as gastropods, polychaete worms.
b) Meiobenthos (Size 0.1 – 0.5 mm) eg. Ciliates, annelids, copepods
c) Microbenthos (size smaller than 0.1 mm) eg. Bacteria, flagellates, ciliates.
ii) Based on mode of nutrients 
a)Phytobenthos : Autotrophic life, include varieties of algae, flowering plants and other
angiosperms, sea grasses etc.
b)Zoobenthos : Heterotrophic life, range from microscopic to macroscopic organisms.
iii) Based on mode of life and area of dwelling
a)Epibenthos : Live on top of the substratum regardless of whether the bottom sediment is soft
or hard rock. eg. benthic molluscs, crabs, prawn, macrophtes, worms, micro algae.
b)Endobenthos : Live within the substratum regardless of its type. Interstitial fauna may be
included under endobenthos, mainly borers and burrowers.
iv)Based on Mobility
a)Sessile : They do not possess any form of mobility but remain attached to the substrate.
eg. Fresh water sponges, macro phytic and microvegetation
b)Vagrant : They possess locomotive powers and can move either rapidly or slowly eg. shelled
macro – invertebrates, worms etc.
v) Based on ooze film
a) Ooze film assemblage : Ooze forming / secreting organisms.
b) Associated ooze film assemblage : organisms associated with ooze film. 
Periphyton
Periphyton are a miscellaneous assemblage of organisms growing upon free surfaces of objects
submerged in water frequently appearing as a brownish green slimy, slippery layer. It commonly
found on plants, wood, stones and various other objects. It seems to develop in littoral and sub-
littoral regions. Periphyton may not always be regarded as true benthos as they tend to grow on
any solid support and likely to contain plankton and benthic organisms
Distribution of benthos
Qualitative distribution 
At different bottom conditions and exposure to wave action diversify the littoral zone and makes
differences in fauna and supports different kinds if animals.
- Depth and quality related distribution
- Decreasing diversity and numbers with increase in depth
- Mostly of uniform distribution
The following animation shows the qualitative depth relations of two American lakes 

The littoral and sub-littoral populations, insects and molluscs comprise as much as 70% of total
number of different components. Increasing depths beyond the littoral zone, the number of
benthic organisms may diminish namely Bryozoans, Platyhelminthes, Snails, Bivalves,
Nematodes, Annelids, insects etc. especially at first 18 m depth. There is no constancy of
composition of different types of lakes because lakes do not have profundal benthic animal
population. Even within a single lake n different composition of population occur at different
places.
The nature of the bottom has selective influence upon the quality of the fauna. Baker (1981)
classified bottom materials in the littoral zone of lake Oneida into 6 different types namely mud,

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sand, clay, gravel, boulders and sandy clay. Microfauna exists in the bottom deposits is also
established and hence microorganisms are considered as bottom dwellers and appear as :
1) free swimming or
2) free swimming only in some stage of life cycle or
3) always spend most of time on the bottom materials
Bigelow (1928) divided littoral benthic microorganisms of lake Nipigon, Canada into two
ecological groups : 
i) the ooze-film assemblage and
ii) the associated ooze–film assemblage 
The former group comprises those microorganisms living in and on that film of ooze at upper
surface of the lake bottom contain certain algae, including diatoms, protozoa, rotifers and
cladocerans, while the latter directly dependent upon the ooze film and swim very close to it eg,
Cladocerans and rotifers . 
9.2. Quantitative and Qualitative movements, seasonal changes and migrations of
benthos
Quantitative distribution 
In shallow lakes, the entire bottom is uniformly productive. The growth of bottom dwellers,
whose quantity of produce at different depths especially in shallow lake, is not same.
The deeper inland lakes can be divided in to 2 groups 1) Those which do not develop chemical
stratification during summer and 2) Those which become chemically stratified and remain so
throughout the summer period.
In non-stratified deeper lakes, the benthic forms extend much deeper into the basin resulting in
greater mass production. However, in stratified deeper lakes during summer, the bottom region
exposed to hypolimnic conditions lead to less production quantitatively than the regions above
them.
Movements and migration of benthos
Under certain circumstances, the benthos exhibits movements within or on the bottom. Certain
other organisms move up and down in streams. Moffett (1943) observed movements of bottom
organisms on large wave swept shoal and found some changes in space due to bottom shifting.
Moon (1935) found that :
(a) the littoral fauna in lake Windermere is in a continual state of movements
(b) this fauna is sensitive to changes is surface level of the lake 
(c) more active elements of the fauna move quickly into the inundate portions of the beach and 
(d) a rise in surface level only 2.5cm is sufficient to produce a movement of the fauna.
Seasonal changes in benthos
Little is known about the seasonal changes in benthic organisms. In very shallow lakes, the
seasonal changes in the whole benthic region are essentially the same as those which prevail in
the littoral zone of deeper lakes. In the abyssal regions of deeper lakes which never overturn
and hence the benthic conditions remain same throughout the year. Benthos are known to show
different morphological changes with respect to different seasons (spring, summer, autumn and
winter) at different zones (littoral, sub-littoral and profundal). In some lakes, the organisms may
compose the overwhelming bulk of entire population and significant fact that each profundal
species manifests similar and corresponding seasonal increase and decrease magnitudes,
which are characteristics of the species.
Origin and permanence of profoundal bottom fauna
The preformed bottom fauna occurs in inland lakes is composed of forms which occur in neither

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peculiar nor restricted to the profundal region, they seem to occupy also the sub-littoral and
some extent the littoral zones. The bottom fauna is not qualitatively unique but is composed of
representative of a few species belonging to larger littoral and sub-littoral population. During
spring and fall overturns, these population spread down the lake bottom into the profundal
region bringing into not only the grater numbers of individuals but certain additional species
characteristics. However, the gradual development of stagnation at the bottom bring about a
condition which some of the species cannot under go. Occasionally in unusually deep lakes,
certain animals occur which never come to shallow depth (above 50m). Hence, it appears that
profundal bottom forms are not truly a permanent profundal fauna but may themselves be
eliminated completely or party by exposing to stagnation.
Vertical distribution of profundal bottom fauna
Evidently it has been assumed that the profundal bottom fauna largely confined to the
uppermost layer of the bottom deposits. Information secured by Lenz (1931) who by using
constructed bottom sampler was able to bring virtually undisturbed, vertical samples of bottom
deposits to the surface and then isolating and examining horizontal strata in their unaltered
relations. Various horizontal levels have been studied down to a depth of 24cm. The largest part
of the fauna occurred in the upper half of the sampler. Some difference appeared in the vertical
distribution of various organisms for eg, Tubificidae concentrated in the upper position, while
Chironomus were distributed from the surface of the mud to a depth of 20cm or more. Moore
(1939) indicated and noted that there was an obvious similarity in the vertical distribution of
micro and macro benthos.

Unit 10- Biological productivity

10.1 Circulation of food material, classification of lakes based on productivity
Circulation of food materials
Essential food stuffs in a lake undergo continuous, more or less definite cycles. Natural water
body is completely closed community since there are various direct and indirect influences
exerted from outside. In considering the circulation of food materials in a typical lake, the
following fundamental facts must be kept in mind.
1)The ultimate basic substances are 
a) inorganic nutritive materials dissolved in water and
b) certain energies and gases from the atmosphere
2)Chlorophyll bearing plankton and flora and certain bacteria can utilize directly basic materials
in constructing living mater
3)Other organisms, plants or animals are of a dependent superstructure
4)Every organisms of a lake population may
(a) by death and disintegration contributes directly to the dissolved materials and detritus or
(b) be consumed as food by other organisms. 
Circulation of food materials
Two general set of processes are construction process and reduction process. Building upward
from the simple food material into the higher is a construction process. Reduction through
bacterial action into simple substances is a reduction process. Cycles are so interrelated that
materials may be built up to the same level in different ways. One cycle through which animals
are freed from dependence upon photosynthetic plants ie from basic dissolved nutritive

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materials through bacteria which change CO2 to organic carbon in the absence of sunlight to
certain animals which consuming them. 

Figure : Circulation of Food material in a Lake

Classification of lakes on the basis of productivity
Many attempts have been made to classify lakes on limnological bases, related in some way or
another to productivity during the past five decades. The combined contributions of a number of
those attempts have been lead to the classification of lakes into three major types viz,
1. Oligotrophic lakes
2. Eutrophic lakes and
3. Dystrophic lakes 

Following Table gives the details of difference between above listed lakes 

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Further, other difference between Oligotrophic and Eutrophic lakes is given below

10.2. Laws of minimum, biotic potential and environmental resistance, quantitative

relations in a standing crop
Law of the minimum
Existence and production of animal and plant life depend upon the proper quantitative and
qualitative composition of the environment for each component organisms. Liebig’s law of the
minimum originally applied to plants and is stated as each organism required a certain number
of food materials and each of these materials must be present in a certain quantity. If one of
these food substances is absent or present in minimal quantity the growth will be minimal. The

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yield of a plant or animal according to this law is determined by the quantity of that particular
substance present in minimal amount as per the demands of the organism.
The law of minimum has had a wide acceptance. Liebig’s law of the minimum is the foundation
of law of limiting factors which is dependent on one factor upon another. Various environmental
factors acted independently wherein if one factor in present in limiting quantity and increase of
other factors would effect no change.
Biotic potential and environmental resistance
The biological productivity of any body of water or any portion of that body of water is the end
result of the interaction of organism present with the surrounding environment. Biotic potential is
the characteristics and abilities inherent within an organism which enable it to exist and
reproduce. It is the sum total of all of those capacities of an organism which determine its
relative success in solving all problems of maintenance. It is a sum of the number of young
produced at each production. 
Every environment contains active features which work toward the control of production in
various organisms involved. Thus the environment resists to a greater or less extent for the
fulfillment of biotic potential. In the long run, nature acts toward a balance between these two
tendencies in which each organism maintains itself in a suitable environment without
overpopulation.
The principles of biotic potential and the environmental resistance together with all of their
associated features are just a pertinent in aquatic biology as in terrestrial situations. Biological
productivity of any aquatic community is a general measure of all of adjustments between biotic
potential and environmental resistance existing within it.
Quantitative relationships in a standing crop
Any body of water maintains a certain standing crop of organisms composed primarily of 5 large
groups viz, phytoplankton, bottom flora, zooplankton and fishes. This series composes a
nutritional chain in which the first two constitute a producing class and the other three are the
consumers. Therefore these organisms are an expression of the productivity of water
concerned.

Figure : Diagram showing weight relationship of Biota and dissolved organic matter 

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The actual values expressed into this pyramid of aquatic life are different to some extent in
different lakes but some form of pyramid is the rule. This figure illustrate the dissolved organic
matter composes approximately 60% of total diagram, the fish only one-half of 1% and the other
animals slightly more than 5%.

10.3. Laws of minimum, biotic potential and environmental resistance, quantitative

relations in a standing crop
Trophic dynamic 
In the trophical dynamics of an ecological system, basic processes are in the nature of transfers
of energy. The ultimate source of such energy is solar radiation. Earlier work have raised the
biological conclusions which seem to have a certain validity like :
a. food cycles rarely have more than five trophic levels
b. separation of an organism from basic source of energy (solar radiation) lead to less chance of
dependence upon trophic level for energy
c. the consumers seem to be progressively more efficient in the utilization of food supply
d. productivity and photosynthesis increase from oligotrophy to eutrophy and then decline in
lake.
Successional phenomena
All environments are dynamic and undergo changes is the fundamental principle of ecology.
These fundamental changes more or less predictable alternations involving expanses of time
and these changes may be due to :
a) the action of predominating inorganic factors in the environment eg, erosion
b) the action of organisms in modifying the environment or
c) the combination of (a) and (b)
One fact common to all situations exists that the various components of the biota must meet the
changing conditions in one of the following ways, such as :
a) adaptation
b) migration and
c) extinction
Ecological successions of various kinds go on in lakes and other inland waters as certainly as
they do on land. The movement of the units of lentic series is in the direction of extinction by
filling of basins and in the lotic series it is in the direction extension of stream length and a
cutting of stream bed to base level. 
Eutrophication
In general and within limits, the productivity increases with the age of a lake. Storm (1928) has
stated the process as follows: 
The natural process of the maturing of a lake is that of eutrophication. The original state of all
lakes must be assumed to be oligotrophic but later due to surplus organic sediment occurring
from life process of a lake is changed to eutrophic condition. The quantities of plankton, oxygen
curves and average depths are the first features to be changed and later the bottom fauna.
It must be clearly understood that the maturing process takes place at very different rates in
different lakes. For example, in northern United States, majority of lakes basin formed during
glacial period but they have matured at different rates and many small basins have long ago
passed the succession stages into old age and are become dry land. If the lake is smaller, a
rapid eutrophication and further extinction take place but certain other lakes fail to go through

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the usual evolution from oligotrophy to eutrophy even though natural filling may render the
hypolimnion smaller than the epilimnion.
Dystrophication
The dystrophic lake basins during their initial stages are low productive (essentially
oligotrophic). These primitive basins varied greatly in size and depth covering considerable
areas having hypolimnion exceeding the epilimnion. During certain circumstances an
incomplete decay of plants and accumulation of humic materials appear the beginning of
dystrophication. After the initiation of dystrophy, the succession progressed by marginal plant
encroachment or by bottom accumulations in incompletely decayed plant materials or by both
and passing through the stages into a peat bog.
Indices of productivity in lakes 
Limnologists have looked for indices of general biological productivity in lakes. There are two
considerations involved and must be kept clearly distinguished, they are (a) the inherent
capacity of a lake to support life (biotic potential) and (b) the actual productivity at a given time.
Obviously one or two indices of productivity would give a dependable evaluation.
1.Average depth : The average depth of a lake is the determining factor for productivity. The
dissolved oxygen content of various layers of a lake is the indicator of richness in nutritive
substances especially at a depth up to 10 m stratum. The dissolved oxygen content is greater in
oligotrophic lakes than that of epilimnion. Certain other features mainly the degree of
development of littoral regions constitutes important influences in determining the production of
the lake. 
2.Rooted submerged vegetation : Kluge (1926) claimed that the amount of rooted, submerged
vegetation may be an index of lake productivity. It is a well known fact that the amount of rooted
submerged vegetation is governed by a number of factors such as degree of exposure and
slope of the submerged shelf. Large lakes for example, lake Nipigon may maintain a great fish
production which could be predicted from the scanty vegetation, whereas the small lakes having
submerged vegetation does not show productivity. 
3.Plankton : Plankton is an index of general production. Eutropic lakes are characterized by
quantitatively rich in plankton, while oligotrophic lakes have a plankton poor in quantity. It has
been claimed that abundance of plankton is associated with rich bottom fauna and paucity of
plankton accompanies a poor bottom fauna.
4.Bottom fauna : European workers have stressed the quality of the bottom fauna at deeper
water as an indication of the productive character of a lake. Form of basin, character of bottom
deposits, water movements etc. would be a true index of the general richness of a lake. When a
rich benthic fauna is present a high total productivity is common. 
5.Organic content of water: The standing crop of dissolved organic matter is much greater than
the total organic matter in the plankton supported by the same water. The dissolved organic
matter is said to be constant in quantity and composition that the character of a lake may be
judged. It has been shown that a constant relation exist between the plankton and the total
organic matter in the water. The presence of organic content has become a new subject to
predict the general index of productivity.
6.Chlorophyll content : Chlorophyll content is used as an index of the photosynthetic capacity.
This measurement can be used as a convenient method for evaluating biological productivity.
7.Other indices : The organic content of bottom deposits is important as food for benthic

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organisms. In general hard water lakes are not highly productive. The total alkalinity and total
phosphorous appear to be the most valuable indices of productivity.
Artificial enrichment
The waste products of human beings and industries often find their way into natural waters and
produce contaminations. Removal of forests and tilling of land bring changes through the
medium of drainage but are not the nature of contaminations. The enrichment of water is due to
the addition of substances and subsequent changes increase the amounts of essential nutritive
materials. Among the contaminations, most likely are the domestic sewages downwash from
manured fields and other organic matter.
The enrichment effects due to contamination are more to be expected in the smaller lakes than
larger ones. Evidences have shown that the sewage from large cites brought into the larger
lakes enters farther into the open lakes and dilutions becomes greater leading to enrichment
having distinctly increased biota.
Sometimes and also at present use of fertilizers have favoured the fish production in the water
bodies. Repeated application of artificial fertilizer to natural waters are dangerous leading to
winter kill of fish in the northern United States lakes. Increasing production in enclosed waters
such as ponds, small lakes and reservoirs is more prominent compared to large natural bodies
and water.

Unit 11- Lotic environments

11.1 Running waters in general
Running waters 
Investigation on running water series is less compared to that of lakes. A comprehensive and
prolonged study has been made in America on Illinois river over a period of 50 years. The lotic
environments differ from lakes and similar waters in the following respect : 
1)Depth – As a rule, the depths of running water units are small compared to lakes 
2)Width of basin - Apart from the channel expansions (sometimes designated as river lakes),
the water is confined to relatively narrow channel.
3)Current - Whole volume of water flows in one direction.
4)Condition of gradient from source to month - All conditions such as physical, chemical and
biological gradually change with distance along the main channel in a definite direction.
5)Extension of channel with age - Stream systems usually increases their length, width and
depth with increasing age.
6)Permanent removal of eroded and transported materials - At any position during the
course of running water unit, materials eroded are transported downstream.
7)Absence of prolonged stagnation - Constant flow and mixing of water usually eliminates
prolonged summer stagnation of the bottom waters.
8)Relative influence of physical factors - Physical factors of the environment are relatively
more important then in lentic waters.
9)Basic food materials - Streams manufacture within themselves the basic food materials but
much depends upon the contribution from surrounding land areas.
Physical and Chemical conditions of Lotic environments
1)Water movement
Current in one direction is the outstanding feature of lotic environments. Current is the greatest
velocity which interfere during water falls and finds minimum where long distances of slope of

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channel becomes negligible during which the channel approaches base level. Some streams
are slow and sluggish throughout their length and other swift throughout their whole course. The
distribution of velocities in streams is determined by different factors such as shape of channel,
roughness of channel, size of channel and slope channel. The maximum velocity is usually
found somewhere within the first one third of the depth of water. The velocity pattern of a stream
altered by strong winds which blowing either sometimes upstream or predominantly
downstream. Ice cover reduces the surface velocity because of the greater retarding effect of
ice as compared with air. The current rate may vary markedly at positions on the front, top,
sides and rear side of submerged stones likewise the same is true of the gaps, channels and
interstices between stones and similar objects. Erosion, transportation and sedimentation are
inseparable accompaniments of stream currents. Those materials which are not carried in
suspension are rolled downstreams. The transporting power of the streams varies with changes
in velocity. 
2)Temperature
In lotic environments, the temperature phenomena are different from those in lentic situations.
The principles features are :
a) tendency toward a uniform temperature at all depths 
b)tendency to follow air temperatures more closely
c)thermal stratification usually absent or temporary 
The temperature variation is common in stream especially in long stream and slow current
situations. Certain circumstances leading to temperature variations are differences in 
a. depth of water 
b. current velocity 
c. bottom materials 
d. temperature of entering of tributary / small stream
e. exposure to direct sunlight 
f. degree of shading and
g. time of day
In long stretches of shallow unshaded, slow moving currents, there is gain in temperature during
the clear days of mid summer, but there is different temperature during night. 
3)Turbidity
A greatest extreme of turbidity occur in the flowing water series. Streams in mountain and rock
beds the turbidity is minimal while in the plain region the turbidity is high (eg, North American
Missouri and Kansas rivers). In some stream systems, high turbidity is a permanent feature
throughout the year. The turbidity in streams is largely due to silt, detritus and other non-living
materials. Since plankton production is commonly restricted, it usually plays a minor role in
turbidity production. Domestic sewage and other forms of stream pollution commonly increase
turbidity.
4)Light
The most important factor in the determination of light punctuation is turbidity and its influence is
great in certain waters which reduce the development of plants. In turbid rivers where light
decreases due to presences of suspended silt in excess of 90% in first 25 mm of water depth.
Chemical conditions
1)Dissolved gases

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Due to the involvement of mechanical condition in water current, the dissolved oxygen supply of
uncontaminated streams is high leading to saturation point. Generally oxygen content begins to
increase soon after sunrise, reaches a maximum shortly after mid-day and then declines to a
minimum. This variation extend from considerable supersaturation to a substantial reduction in
some instances is due to oxygen production by green plants during day and consumption by
respiration of biota and decay of organic matter during both days and night. Under some
circumstances, some rivers show very distinct diurnal oxygen pulses. The factors affecting the
dissolved oxygen are: character of stream flow, slope of channel, temperature, oxygen released
by chlorophyll bearing plants, oxygen consumed in respiration of the biota and oxygen
consumed in the decay of organic deposits on the bottom.
2)Dissolved solids
The dissolved solids vary greatly depending upon the regional characteristics of the drainage
basin. It is known that in general the lotic water contains more salts and less soluble nitrogen
than lentic waters (Chapman, 1931). The dissolved solid content is often subjected to changes
by dilution or addition at stream junctions. Loads of solids in streams are common and it has
been estimated that the range of total solid in the river systems of Upper Peninsula is 100 to
200 ppm while in Lower Peninsula is 200 to 500 ppm.
3)Hydrogen ion concentration
The general features of pH in lotic environment are not different from that of lentic
environments. Currents tend to keep the pH uniform over considerable distance; it keeps any
acidity due to accumulating free carbon dioxide reduced. However, a uniform pH over
considerable distance in lotic environment is kept. So, the streams would never develop more
intense acidities unless they are contaminated or receive heavy seepages from mineral
deposits.

11.2 Biological conditions, productivity features of lotic environments, Influence of

currents,Plant growths
Biological conditions 
The range of conditions is reflected in the biota which varies from distinctly characteristic
organisms to lentic flora and fauna of the system. The greater the current velocity, the greater
will be the divergence of lentic and lotic populations. The lotic assemblages occupy the upper
elevated positions of stream system. Certain groups of animal exclusively lotic ones such a
stone–fly larvae (Plecoptera), black-fly larvae (Simuliidae) etc. are restricted to running water
series. Generally it is said that the lotic populations are more restricted in the number of
species.
Productivity features of lotic environments
The quantitative features of plankton and productivity of the microorganisms are very important
in the lotic series of water bodies. According to Needham (1930), the ecology of the smaller
streams especially brooks and creeks have general features and they are :
1. Distribution of aquatic animals in brooks and creeks are dependent largely upon
a) temperature of the water
b) nature of the bottom
c) velocity of current
2. Smaller streams from the source of mouth have 2 distinct types of habitats such as pools and
riffles
3. Riffle bottoms greatly exceed the pool bottom in productivity 
4. Fishes in brooks and creeks tend to seek the pools. Pools act as catch basins for animals

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brought down from the riffles and these drift animals serve as food for pool fishes.
5. Absences of higher aquatic vegetation leading to productive silt bottom for small organisms
6. Plant beds in smaller streams markedly affect productivity. Stream bottom supporting growths
of aquatic plants were found seven times more productive than stream bottoms bare of
vegetation.
7. Small cold, headwater streams less than 7 ft are more productive (twice) than the maximum
width.
Influence of current
Lotic fauna is typically composed of animals whose dissolved oxygen demand is fulfilled by the
oxygenated waters of streams. In some forms, the requirement of inherent current demand is
more when compared to oxygen demand in lotic environment. The gill area of May fly nymphs in
mountain streams varied inversely with the dissolved oxygen content, but not close relation with
the current rate. Some organisms are more dependent upon the increased amount of dissolved
oxygen and the mineral salts in solution than upon current. In certain May fly nymphs and
caddis – fly larvae from a swift stream, the oxygen consumption is higher than the closely
related , equal sized nymphs and caddis – fly larvae from a pond. The same situation is exit in
aquatic crustaceans (Asellus aquaticus), where swift stream consume more oxygen than that of
sluggish flowing water. Nymphs are less resistance to oxygen deficiency than those in quiet
water.
Running water animals seek certain physico-chemical conditions and are compelled to tolerate
current as a mechanical condition against which they struggle. With the exception of the
plankton, all other biota have developed means of maintaining themselves (except during
floods) in the streams have adopted to thrive well and even exist. Among organisms of running
waters, the character facilitating to maintain their position is diversified. Some of the insect
larvae build heavier cases in running water than the quiet water.
An accompanying feature of current is molar action. The action and severity of this vary widely
with current velocity, the nature of the bottom materials and other circumstances. Injury and
mortality may be very high at times of flood. In addition to molar action, eroding and scouring
action of flood waters often depopulate the streams, so that restocking is necessary.
Nature has numerous and effective means of restocking populations such as upstream or down
stream migration of animals from adjacent waters, transportation by currents, reproduction by
the individuals, spread of aerial adults from nearby waters and transportation as wind -blown
materials. The production of drift materials in a stream is one of the invariable effects of the
current, even in the absence of floods.
A phenomenon arising out of current action is the depositions of eroded silt. In some lotic
conditions, e.g. Instep, rock bed streams such deposits are absent. Certain organisms regularly
occupy regions of silt deposit with no detrimental effects. Some of the organisms possess
certain mechanical means of avoiding suffocation by the mud. Another influence of current is
the elimination of surface breathers except in the very slow moving regions.
Plant growth
The larger aquatic plants do not occur in stream except where the current is greatly reduced.
Occasionally, certain rooted plants (Chara and others) may be found in rapid streams. There is
distinct tendency to dwarfing in plants that grow in mountain streams. Certain water mosses are
found in rapid current, especially in streams not subject to severe floods. Sometimes, it is also
important to note that slow moving sluggish streams become plant choked maintaining flora
composed of submerged and floating types of plants. In the slower water of stream edges, a
narrow margin of aquatic plants occur but with limited success. The plants provide benefits in
fish production owing to supply shelter, protection for young ones, abundance of food; aquatic
invertebrates which are sometimes become food for fishes.
Algae of swift streams possess the holdfast cells or other structures to support over any
substratum and to remain there against the strong current. The fresh water red algae, Lamanea

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is found only in rapid waterfalls. Green algae, Cladophora, attaches to stones and other
supports in slow streams. Certain other algae which lack holdfasts resist current action because
of abundant mucous secretion.
11.3 Nekton, benthos, Temporary and head water streams, general ecological succession
Nekton – benthos characteristics 
Since streams are shallow water bodies and hence they do not support organisms inhabiting at
the bottom but very rarely the deep waters of rivers support such organisms. In most of the
streams a sharp dividing line between nekton and benthos on natural basis does not exist. Of
the vast communities of organisms involved, fishes are the only group which might be called
nektonic. The benthos and nekton have been jointly grouped to have abundant quantities in
shallow moving water bodies.
Temporary and head water streams
During dry seasons, the channels are either completely devoid of water or at most contain few
isolated pools. Such water may not support life but really and especially during certain season
certain assemblages of organisms are present which possess the following features :
1. Life histories requiring water only in a portion of the cycle and therefore such streams do not
support aquatic residents.
2. No development of higher aquatic plants but few plant eaters available among the fauna.
3. Very rarely found carnivorous who feed upon detritus and microorganisms.
4. Positive rheotropism in some of the motile forms: cling or attach to the substrates.
5. Some of the tolerant species may carry over in pools from one flowing water system to the
other. 
6. Linear sequence distribution of fishes. With the onset of drought, some fishes move
downstream, with rise of water level, they move towards upstream but maintaining their general
distribution.
7. Aquatic insects form the most diversified group of the fauna
The two large types of stream communities are a) those characteristics of swift waters with
hard, stable bottom and b) those characteristics of slow moving waters with soft, unstable
bottoms. In swift water type, there is no pelagic community. The sluggish water type depends
upon soft bottom and slow current, decaying organic matter, accumulated on bottom, support
large quantities of bacteria, motile pelagic communities and remain in no fixed position in
relation to bottom.
General ecological succession
It is a known physiographic fact that the head water region of running water unit / system will
migrate. Streams for example extend their growth and proceeds by continually cutting back at
their back (source). As the time goes on, the young stream condition migrate upstream with the
migration of head waters and older set of environmental conditions move upstream to occupy
the level occupied by young stream environment.
The migration of stream dependent largely upon the rate of erosion and transportation of
materials at the source and is a slow process. An interesting development in the subject of
succession of stream communities as pointed out by Shelford and Eddy (1929) that permanent
stream communities exist, undergo successional development, reach and maintain a quasi–
stable condition and manifest seasonal and annual differences as do terrestrial and marine
communities.

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