GENERAL ECOLOGY OF LENTIC SYSTEMS

Lakes, both large and small, are characterized by great heterogeneity in physical factors. Abiotic
factors vary greatly among locations and depths within a lake. This variation has profound effects
on the plants and animals in the system and plays a key role in their distribution and abundance in
the lake.
Light: As in any ecological system, the patterns of light intensity play a crucial role by virtue of
their influence on the distribution of plants and algae. Water absorbs sunlight, so the amount of
solar input available to plants decreases with depth. Any suspended material in the water may also
absorb or scatter light and thus decrease the amount available at greater depths. The upper layer of
a lake, the layer with enough light for photosynthesis, is called the trophogenic zone. Below this
region is the tropholytic zone, were light intensities are insufficient to support plants. At certain
depth, called the compensation point, the rate of photosynthesis by plants exactly balances the rate
of respiration by plants and animals. Photosynthesis balances respiration when the absorption of
sunlight has reduced the light available to approximately 1% of the sunlight reaching the surface
of the lake. The exact depth of the compensation point varies with amount of incident sunlight, the
turbidity of the water, and the rate of respiration.
Temperature also changes with depth in lentic systems because of the relationship between
density and temperature. Like most substances, water becomes denser as it cools. However, it
reaches the maximum density at 4oc. below this temperature, its density decreases again. This is
why ice cubes float in a drink and why lakes freeze from the surface down. Waters of different
temperature, and thus different density, do not mix readily. As a result, the water in lakes becomes
stratified by temperature. In deep lakes, the degree of stratification and mixing undergoes seasonal
changes as a result of the pattern of temperature changes in air.
The thermal stratification of lakes affects other physical factors as well.
Oxygen: There may be a strong oxygen concentration gradient in the lake associated with the
summer temperature stratification. Three processes determine the oxygen content of water in a
lake.
1. Oxygen enters the lake at the surface via exchange with the air;
2. The activity of photosynthetic algae and other organisms add oxygen to the water
3. Respiration by animals, plants and microorganisms remove oxygen from the water.
In the summer, oxygen concentrations are high in the epilimnion as a consequence of exchange
and photosynthesis. At the same time, however, dead and decaying, materials sink into
hypolimnion, where it is decomposed by bacteria. Here, oxygen may be greatly depleted,
especially because this lower layer doesn’t mix with the oxygen rich epilimnion. Therefore, the
hypolimnion typically has low oxygen concentrations during the summer.
Nutrients may also be stratified as a result of these processes. As algae photosynthesize, they use
the nutrients in the epilimnion. Those nutrients are removed from circulation. In the fall, as the
surface of the lake cools, the temperature becomes more uniform, and mixing occurs. This is called
the fall turnover. During the winter, the temperature of the lake is uniform, and a layer of ice may
form on the surface. Because of the ice, winds do not circulate the water. When ice melts in the
spring, the surface waters warm, and spring winds mix the water again.
Organisms: In addition to the zonation in a lake’s physical parameters, there is zonation in terms
of the organisms that inhabit the lake. Near the shore, light can penetrate to the bottom, and rooted
plants can survive. The region is referred to as the littoral zone. Water too deep for rooted plants
is inhabited by plankton, tiny free floating or swimming organisms in the pelagic region. The
waters of a lake beneath the littoral zone and above the compensation point make up the limnetic
zone. In many lakes, the density of phytoplankton is stratified by depth by virtue of both physical
processes (buoyancy characteristics) and active movement to regions of optimal growth condition.
Below the compensation point, photosynthesis is greatly diminished. The organisms that live here
are generally not green plants but animals or microorganisms that depends primary on the rain of
organic matter (dead organisms) from above as an energy source. This region is the profundal
zone. Finally, the bottom layer of lake in both the littoral and the profundal zones is the benthic
zone.
Lake systems can be broadly classified as oligotrophic or eutrophic on the basis of their nutrients
content and rate of photosynthesis. Oligotrophic lakes are characterized by low nutrient content,
especially of phosphorous. They have small populations of photosynthetic algae and hence low
rate of photosynthesis. The low densities of algae result in less material for decomposition in the
hypolimnion and hence higher oxygen concentrations there. Oligotrophic lakes are often young
and have not yet accumulated sufficient nutrients from the land to support high levels of
photosynthesis. They also frequently have low surface –to volume ratios; this results in relatively
low solar input to support photosynthesis.
Eutrophic lakes are characterized by high levels of nutrients and high rates of photosynthesis.
Large populations of algae develop and provide a steady supply of organic material for
decomposing in the profundal zone. Thus the hypolimnion may become oxygen depleted in such
lakes. Indeed, during the summer months, oxygen depletion may be so severe that it causes the
death of bottom- dwelling fish. Although eutrophication of a lake can occur through natural
process as a lake ages and accumulates nutrients, the process is accelerated when humans greatly
increase the nutrient content. This often happens as a result of fertilizer runoff from agricultural
land.

Lotic Systems
Flowing water ecosystems vary in structure and types of habitats and physical factors play a crucial
role in the ecology of flowing water. Among the most important of these is current. The rate of
flow is determined by the stage of the streambed, the volume of water in the stream, and the nature
of the resistance downstream. The volume of water in a stream and river can change dramatically
over a very short period of time. Storms or rapid snowmelt can greatly increase the rate of flow. If
a stream encounters resistance, it does when it enters a lake, flow decreases near the point of
resistance. The density and diversity of life –forms in the water and on the bottom are determined
largely by the flow rate.
Fast flowing streams and rivers have rocky bottoms because smaller particles of dirt and silt are
carried away. Of course, within a stream the current speed changes as the water enters riffles and
ponds and the nature of the bottom substrate changes accordingly. Slow moving streams are more
likely to have muddy bottoms.
Most rivers and streams are too shallow to have temperature stratification, but variations can occur
along the length of a lotic system. Narrow, deep portions of stream are cooler than broad, shallow
portions. The amount of shading from streamside vegetation also affects water temperature. In
addition, moving water is less likely to freeze during winter.
The oxygen content of lotic systems is generally higher than in lentic systems. A greater surface –
to-volume ratio in streams and rivers than in lakes, as well as aeration from churning, accounts for
this.
Streams differ fundamentally from lentic systems in the nature of their energy source. Most of the
energy for stream ecosystems comes from outside sources (allochthonous sources). Streams
receives large amount of litter and detritus from the stream basin, and for most streams this source
provides a far larger proportion of the energy available to the system than that derived in situ by
photosynthesis (autochthonous source).
In lotic systems, most of the photosynthesis is done by plants attached to the substrate. These plants
known as periphyton, reach their highest density in riffles where the water is moving rapidly over
rocks. They are susceptible to scouring, however, if the current increases rapidly, such as after a
storm. Stream periphytons generally have very high reproductive rates and can recolonize and
grow quickly after scouring.
The bulk of the primary consumers in a stream are detritivores that obtain energy from the
allochthonous inputs.
Stream ecosystem ecology changes markedly over the course of its length, a concept known as the
river continuum. Near the source of the river, autotrophic production is limited (often by shade
from terrestrial vegetation) such that respiration exceeds production. Much of the organic input to
the stream is from allochthonous sources. Consequently, much stream heterotrophs are adapted to
use such materials – many are detritivores. Downstream, the relative importance of terrestrial
vegetation decreases in the wider stream. Accordingly, temperature increases. Higher levels of in
situ primary production may exceed respiration. The wider stream, with open swifter water as well
as near-shore eddies and pools, has greater variety of micro habitats and this gives a greater total
species diversity. As the size of the system continues to increase, another ecological shift occurs.
The current slows and sediments accumulate, and production declines again.

Wetlands
At the interface between freshwater and terrestrial systems are a number of semiaquatic habitats
collectively term wetlands. These are unique environments from ecotones between terrestrial and
adjacent aquatic ecosystems, sharing characteristics of both. Wetlands, habitats that are perpetually
or periodically flooded, are of crucial importance to a number of vertebrates and invertebrate
organisms as breeding grounds, wintering areas and feeding sites. Wetlands are found in every
climatic zone but are local in occurrence. Wetlands support specialized plants that occur where the
soil condition remains saturated for most or all of the years. These hydrophytics (water loving
plants) are adapted to grow in water or on soil that is periodically anaerobic because of excess
water. Hydrophytic plants are typically classified into one of three groups
1. Obligate wetland plants that require saturated soils. Example are the submerged pond
weeds, floating pond lily, emergent cattails and bulrushes, and trees such as bald cypress.
 2. Facultative wetland plats that can grow in either saturated or upland soil and rarely grow
elsewhere, such as certain edges and alders and trees such as red maple and cottonwoods
3. Occasional wetland plants that are usually found out of wetland environments but can
tolerate wetlands. It is the last group of plants that is critical in determining the upper limit
of a wetland along a gradient of soil moisture.
We distinguished a number of different types of wetlands on the basis of topographic situations
the nature of the drainage. These wetland types sometimes grade into one another, making
precise definitions difficult.
Basin wetlands develop in shallow basins ranging from upland depression to fill in lakes and
ponds. Water flow is vertical, involving precipitation and the downward infiltration water into
the soil.
Riverine wetlands develop along shallow and periodically flooded banks of rivers and
streams. What flow is unidirectional?
Fringe wetlands occurs along the coast of large lakes. Water flow is in 2 directions, because
it involves rising lake level and tidal action.
Humans often covet these lands for various kinds of development, and thus only a tiny fraction of
the original wetlands of the United States remains.

Marshes
These are wetlands dominated by emergent soft stem herbaceous vegetation. These are essentially
wet grasslands. The most conspicuous marsh plants are emergent, plants that are rooted in the
substrate but protrude above the shallow water. Other members of the communities such as pond
lilies float on the surface. Marshes originate in a number of ways; they may be formed when lakes
or ponds fill in with vegetation. The edges of lakes or rivers were water flow is reduced may also
develop marsh vegetation. Marshes may represent spatial extensive, continuous systems or small,
discrete wetlands.

Swamps
These are forested wetlands that occur in flat basins through which water flows slowly, saturating
the soil. They differ from marshes in that they are dominated by woody plants. The species that
dominate swamps differ regionally. The vegetation in a swamp can vary markedly with the depth
of standing water.

MARINE SYSTEMS
Ecologists divide marine ecosystems into two broad categories, coastal and open water systems.
Marine biology is the scientific study of the organisms that live in the sea. The ocean is a vast
realm, home to countless strange and wonderful creatures. It is often the beauty, mystery, and
variety of life in the sea that attract students to a course in marine biology. Even professional
marine biologist feels a sense of adventure and wonder in their studies.
Marine biology is not really a separate science but, rather the more general science of biology
applied to the sea. Nearly all the disciplines of biology are represented in marine and freshwater
biology. There are marine and freshwater biologists who study the basic chemistry of living things,
for example. Others are interested in whole organisms: the way they behave, where they live and
why, and so on. Other marine biologists adopt a global perspective and look at the way entire
functions function as systems.

The Physical/Chemical Nature of Oceans

The Waters of the Ocean
Marine organisms are mostly water 80% or more by weight in most cases, and in jellyfishes over
95%. Water not only fills oceans, it makes life itself possible.

The Unique Nature of Water

In all other substances, two or more atoms are chemically combined into larger particles called
molecules. Water molecules are made up of one oxygen atom, which is relatively large and two
small hydrogen atoms. The oxygen and hydrogen atoms have weak, opposite charges that create
electrical attractions, or hydrogen bonds, between adjacent water molecules. Hydrogen bonds are
weaker than the covalent bonds that hold the water molecule together, but they make water
different from any other substance on earth.

The Three States of Water

Any substance can exist in three different states, or phases: solid, liquid, or gas. Water is the only
substance that naturally occurs in all three states on earth.

Heat and Water

Water has the highest latent heat of melting and highest latent heat of evaporation and one of the
highest heat capacities of any natural substance. Not only does ice melts at a comparatively high
temperature, it absorbs a lot of heat when it melts. The amount heat required to melt a substance
is called latent heat of melting. It thus takes a long period of very cold weather before a body of
water will freeze.

Water as a Solvent
Water can dissolve more things than any other natural substance and is called a universal solvent.

Sea Water
The characteristic of seawater are due both to the nature of pure water and to the materials
dissolved in it. Some of the solids dissolved in seawater are produced by the chemical weathering
of rocks on land and are carried to the sea by rivers. Other materials come from the earth’s interior.
Most of these are released into the ocean at hydrothermal vents. Some are released into the
atmosphere from volcanoes and enter the ocean in rain and snow.

Salt Composition
Seawater contains at least a little of everything but most of the solutes or dissolved materials are
made up of a surprisingly small group of ions. In fact, only six ions composed over 99% of solids
dissolve in sea water.

Sodium and chlorine account for about 85%; this is why seawater tastes like table salt. When
seawater evaporates, the ions in it are left behind and combine to form various salts. Salinity is
defined as the total amount of salt dissolved in seawater. It is usually expressed as the number of
grams of salt left behind when 1000grams of seawater are evaporated. For example if 35grams of
salt are left from evaporating 1000grams of seawater, then the sea had a salinity of 35parts per
thousand or 35‰ .
The chemist William Ditmar analyzed seawater samples from the challenger expedition and found
that the percentages of the major ions in seawater remain constant even though the total amount of
salt varied slightly from place to place. The principle is called the rule of constant proportion which
states that “the relative amounts of the various ions in seawater are always the same”. Thus
although marine organisms are exposed to changes in total salinity; they rarely have to deal with
changes in the ratios of the various ions. This makes it easier for them to control their internal salt
and water balance.
The average salinity of ocean is about 35‰. The open ocean varies relatively little between about
33‰ and 37‰, depending mostly on the balance between evaporation and precipitation. Partially
enclosed seas may have more extreme salinities. The red sea for instance is in a hot dry region
where evaporation pre-dominates over precipitation, so it is very salty, about 40‰. Near coast or
in enclosed basins, runoff from rivers may have a strong effect. River runoff gives the Baltic Sea
a typical salinity at the surface of only about 7‰, for example.
The other dissolve materials include substances such as essential nutrients that, though present in
any tiny amounts, are critically important to marine life. Compounds containing nitrogen,
phosphorus, and iron are particularly important and their availability determines the biological
productivity of much of the oceans. Primary production by photosynthesis requires nutrients as
well as light. Nitrogen, phosphorus, silica and iron are the most important nutrients in the sea.

Salinity, Temperature and Density

Salinity and temperature of sea water determines its density. It gets denser, saltier, colder, or both.
The saltier the water the denser it is. Temperature in the ocean varies between about -2o and +30OC.
Temperatures below 0OC are possible because salt water freezes at a colder temperature than pure
water. Temperature in the ocean varies considerably more than salinity, so as a practical matter
density is usually controlled more by temperature than salinity. Exceptions do occur, however and
both the temperature and salinity of ocean water still needs to be measured to determine the
density. Conductivity- temperature-depth meters (CTDS) are widely used in conjunction with
other instruments. Today, oceanographers increasingly make their measurements with automated
instruments since to study a large area, the ships has to move from place to place. Another way to
get the big picture is to use satellites. Satellites can measure conditions only near the surface, but
they give instantaneous coverage of a large area. Furthermore, a series of measurements can be
made in a short time. This makes it possible to follow rapid changes in surface conditions that
result from currents and weather.

Dissolved Gases
There are gases as well as solid materials dissolved in seawater. For organisms the most important
gases in the ocean are oxygen, carbon dioxide, and nitrogen. All three are found in the atmospheres
and dissolve in seawater at the sea surface. Sometimes the reverse occurs and the sea surface
releases gases to the atmosphere. This process is known as gas exchange between the ocean and
atmospheres. Unlike solids, gases dissolve better in cold than in warm water. So dissolved gas
concentrations are higher in polar waters than in the tropics. The amount of oxygen in the water is
also strongly affected by organisms through photosynthesis and respiration. Much of the oxygen
produced by photosynthesis in the ocean is released to the atmosphere. CO2 makes up more than
80% of the dissolved gas in the ocean, and the ocean stores more than 50times as much total CO2,
as the atmosphere

Transparency
One of the most biologically important properties of seawater is that it is relatively transparent, so
sunlight can penetrate into the ocean. This is vital because all photosynthetic organisms need light
to grow.

Pressure
Another factor that changes dramatically with depth in the ocean is pressure. Organisms on land
are under 1 atmosphere (psi) of pressure at sea level- the weight of all the air above them. Marine
organisms, however, are under the weight of the water as well as the atmosphere. Since water is
much heavier than air, marine organisms are under much more pressure than those on land. The
pressure increases dramatically with depth because the amount of water above gets greater. With
each 10m of increased depth, another atmosphere of pressure is added. As the pressure increases
gases are compressed. Gas-filled structures inside organisms like air bladders, floats and lungs
shrink or collapse. This limits the depth range of many marine organisms. It also means that
submarines and housings for scientific instruments must be specially engineered to withstand
pressure. This greatly increases the difficulty, expense, and sometimes the danger of studying the
sea. The reverse effect also causes problems: organisms that contain gas-filled structures may be
injured when brought up from the deep.

THE 3-LAYERED OCEAN

The ocean has 3 principle layers the surface layer is usually 100m to 200m deep. The surface layer
is mixed by wind, waves, and currents, so it is also known as the mixed layer. The intermediate
layer lies below the surface layer, typically 1000 to 1500m depth. The main thermocline, a
transition zone between warm surface and cold water below, lies in the intermediate layer. Below
about 1,500m lie the deep and bottom layers. Technically, deep water and bottom water are
different, but they are similar in being uniformly cold, typically less than 4oc.

Stability and overturn

Most of the time surface water, being warmer and less dense, floats on top of the denser water
below unless wind or wave energy stirs up the water column. Such a water column is said to be
stable. How stable the water column is depends on the density difference between the layers. When
the surface layer is only slightly less dense than the layer below, less energy is required to mix the
2-layers and the water column has low stability. A highly stable water column, on the other hand,
results from a large density difference between deep and shallow water, and requires much more
energy to mix.
Sometimes water columns become unstable, meaning the surface water becomes denser than the
water below. The surface water sinks, which is called down welling, and displaces and mixes with
deeper water. This process is known as overturn. Because surface water, all with the same
temperature and density, is descending through the water column, the temperature and density
profiles are vertical straight lines, and oceanographers identify conditions of overturn by looking
for such straight-line profiles. Overturn usually occurs in temperate and Polar Regions during the
winter when the surface water cools.
When the overturn occurs over only a small area or when the surface water is only slightly denser
than the water below, the down welling water simply mixes with deeper water, so that the mixed
layer extends deeper in the water column. This mixing is important for the productivity of
temperate and polar waters. When down welling is intense, however a large volume of water may
have the surface without mixing much with surrounding water.

WAVES AND TIDES

Waves and tides are among the most visible of all ocean phenomena. Anyone who has swum in,
sailed on, or simply walked beside the sea is familiar with waves and tides.
Waves
The wind not only drives surface currents, it causes waves. The highest part of a wave is called the
crests and the lowest part the trough. The size of an ocean wave is usually expressed as the wave
height, which is the vertical distance between the trough and crest. Wave crests or troughs can be
together or far apart. The distance between them is called the wavelength. The time a wave takes
to go by any given point is called the period of the wave.
When under a wave crest the water moves up and forward; under the troughs it moves down and
back. On the whole the water particles don’t go anywhere at all as the wave passes; they just move
in circles. Though waves carry energy across the sea surface, they do not actually transport water.
Waves begin to form as soon as the wind starts to blow. The faster and longer the wind blows, the
larger the waves get. The size of the waves generated by the wind also depends on the fetch, the
span of open water very which the wind blows.
While the wind is blowing it pushes the wave crests up into sharp peaks and stretches out the
troughs. Waves like this are called seas. The waves move away from where they are generated
slightly faster than the speed of the wind. Once away from the wind the waves settle into swells.
With their smoothly rounded crests and troughs, swells are very similar to the ideal waves shown
in. when the waves approach shore and get into shallow water, they begin to feel the bottom. The
bottom forces the water particles to move in elongated ellipse instead of circles, which slows the
wave. As the waves behind catch up the waves get closer together, giving them a shorter
wavelength. The waves pile up, becoming higher and steeper. Eventually they become so high and
so steep that they fall forward and break, creating surf. The energy that was put into the wave by
the wind is expended on the shoreline as the wave breaks.
The sea surface is usually a confused jumble instead of a series of nice, regular waves moving in
one direction. This is because the surface at any given location is affected by a mixture of waves
coming from many different places, generated by winds of different speeds blowing in different
directions for different lengths of time. The complex surface of the ocean results from the
interactions of all these waves.

Tides
The tides are a dominant influence on near shore sea life. They alternately expose and submerge
organisms on the shore, drive the circulation of bays and estuaries, trigger spawning and influence
the lives of marine organisms in countless other ways.
Tides are cause by gravitational pull of the moon and the sun and by the rotation of the earth, moon
and sun. The moon and earth are held together by gravitational attraction. The gravity is strongest
on the side of the earth closest to the moon. Here the moon’s gravity pulls the water in the ocean
toward the moon, so that if the earth were covered entirely by water the ocean would bulge towards
the moon. On the opposite side of the earth further away from the moon, the moons gravitational
pull is weaker, so the water does not move towards the moon. In fact, it bulges in the opposite
direction, away from the moon, because of centrifugal force. This centrifugal force arises because,
strictly speaking, the moon does not revolve around the earth. Instead, the earth and moon both
rotate around their combined center of mass, which lies inside the earth but offset slightly from
the actual center of the earth. This offset causes the earth-moon system to wobble slightly, like an
unbalance tire, and creates are centrifugal force that pushes away from the moon. Thus, on a water
covered earth, the water would form 2 bulges on opposite sides of the planet, one bulge towards
the moon where the moons gravity predominates and the other bulge away from the moon where
the centrifugal force predominates. The water will relatively deep under the bulges and shallow
away from the bulges.
In addition to the rotation of the earth, and moon, the earth is spinning like a top in its own axis.
As it does so, any given point on the planet’s surface under a bulge and away from it. High tide
occurs when the point is under a bulge. Because the earth takes 24 hours to complete a rotation,
the point will have 2 high tides and two low tides every day. Actually the moon advances a little
in its own orbit in the course of 24 hours. It takes the point on earth an extra 50 minutes to catch
up and come directly in line with the moon again. A full tidal cycle therefore takes 24 hours and
50 mins.
The sun produces tidal bulges in the same way as the moon. Though the sun is much larger than
the moon, it is 400 times farther away, so the effect of the sun on the tides is only about half as
strong as the moons. When the sun and the moon are in line with each other, which happens at the
full and new moon their effects add together. At these times the tidal range, or difference in water
level between successive high and low tides, is large. Such tides are called spring tides because
they seem to surge up like a spring of water. The name has nothing to do with the season; spring
tides occur throughout the year.
When the sun and the moon are at right angles their effects partially cancel each other. During
these neap tides, the tidal range is small. Neap tides occur when the moon is in the first and third
quarters.
Tides vary from time to time and place to place because of the effects of continents, islands and
bottom topography.