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 1. Details of Module and its Structure

Module Detail

Subject Name <Ecosystem Ecology>

Paper Name <Ecology>
Module Name/Title <Ecosystem Function>
Module Id <Module Id>
Pre-requisites <Basic knowledge about ecosystem, its living and non-living
components and ecosystem structure>
Objectives <To appraise the inter-relations and functional relationships
existing between living and non-living components of the
ecosystem>
Keywords <ecosystem>, <ecosystem function>, <energy flow>, <mineral
cycling>

Structure of Module / Syllabus of a module (Define Topic / Sub-topic of module)

<Ecosystem> <Ecosystem function>, <energy flow>, <food chain and food web>,
<ecological pyramids>, <mineral cycling>, <productivity>

2. Development Team

Role Name Affiliation

Subject Coordinator <Prof. Sujata Bhargava> Savitribai Phule

Pune University
Paper Coordinator <N.S.R. Krishnayya> MS University
Baroda
Content Writer/Author (CW) < Dr. Gyan Prakash Sharma Delhi University
& Neha Goyal>
Content Reviewer (CR) <N.S.R Krishnayya>
Language Editor (LE) <N.S.R Krishnayya>

Ecosystem Ecology
Ecology
Ecosystem Function
 TABLE OF CONTENTS (for textual content)

1. Ecosystem function
2. Energy flow
3. Food chain and food web
4. Ecological pyramids
5. Mineral cycling and productivity

Introduction

The way we study, the external and internal morphology and physiology for understanding the
structure and functioning of an organism and its life processes, similarly we can understand the
ecosystem by studying the structural and functional attributes of all its living (biotic) and non-living
(abiotic) components. Closely-linked structural and functional attributes are critical for the
continued operation of an ecosystem and their integrated understanding provides a complete
conceptualization of the ecosystem dynamics. An ecosystem has a function; each component plays
a specific function while summed up functions of all components defines ecosystem function on a
whole. Importantly, function means showing activity, and does not imply that organisms perform
purposeful roles in ecosystem-level processes. Ecosystem functioning reflects the collective life
activities of plants, animals, and microbes and the effects these activities (feeding, growing,
moving, excreting waste, etc.) have on the physical and chemical conditions of their environment. A
functioning ecosystem exhibits biological and chemical activities characteristic for its type. For
instance, a functioning forest ecosystem exhibits rates of plant production, carbon storage, and
nutrient cycling that are characteristic of most forests. If the forest is modified to an agroecosystem,
its functioning changes. The concept of “ecosystem function” is based on a perspective that focuses
on the whole system and the functional roles played by each component within it. However, the
normative concept of ecosystem functioning also extends towards societal interests. Giller et al.
(2004) characterize ecosystem functioning by three categories of ecosystem functions, namely,
ecosystem processes, ecosystem properties, and ecosystem values (composed of ecosystem goods
and services). To exemplify, ecosystem function of a lake ecosystem refers to processes and the

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 causal relations that give rise to an ecological system, the role of organisms within it, the
interactions between organisms of different species, overall processes that sustain it, long-term
dynamics of different populations, overall biomass of the system, flow of energy and nutrients
within it, and finally to the services it provides to human beings or other organisms.

Energy flow

In an ecosystem, there is a continuous interaction between plants, animals, and their environment to
produce and exchange materials. The energy needed for this cycling of materials comes from the
sun. Green plants, or producers, or autotrophs fix the solar energy into chemical energy. In this
stored form, other organisms take the energy and pass it on further to other organisms. During this
process, a reasonable proportion of energy is lost out of the living system. The whole process is
called flow of energy. An ecosystem may be comprehended of any dimension, where the living and
non-living systems are involved in a continuous flow of energy. The functional processes in an
ecosystem are closely related to its structure and vice-versa. For instance, the flow of energy is
based on trophic or food and feed structure of the ecosystem (producers- plants; consumers-
herbivores, carnivores, and omnivores; decomposers- microbes). Energy flows from sun to primary
producers, to herbivores and then to different levels of carnivores are generally termed as trophic
levels/trophic structure of the ecosystem. The flow of energy is unidirectional and non-cyclic
(Figure 1).

Ecosystem Ecology
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Ecosystem Function
 Figure 1. A schematic representation of energy and nutrient movement in ecosystems. Energy flow
is unidirectional and non-cyclic, whereas nutrient movement is cyclic

An energy flow model or energy flow diagram is a representation of trophic levels linked with each
other showing the inputs and losses of energy incurred during energy transfer at each trophic level
(Figure 2). Lindeman firstly proposed such a model in 1942 assuming that plants and animals can
be arranged into trophic levels; and laws of thermodynamics hold for energy transfer from one
trophic level to the other.

Figure 2. An expanded three link energy food chain (1. Plants, 2. Herbivores, 3. Carnivores)
illustrating the relationship between food flow diagrams and energy transfer. The transfer of energy
becomes degraded, dispersed, and diminished as the energy within a food chain flows from one
trophic level to the other. Abbreviations: I=input, A=assimilation, R=respiration, NU=not utilized,
P=production, B=biomass
(Source: [Link]

According to Raymond Lindeman (1942) “ten percent law”- during the transfer of energy from one
trophic level/structure to another trophic level, only about ten percent of the energy is utilized. The

Ecosystem Ecology
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Ecosystem Function
 remaining energy is lost during transfer, broken down in respiration, or lost to incomplete digestion
by higher trophic levels.
In a “universal” model of energy partitioning in the individual or species population, living
structure or biomass is represented as a box (Figure 3). Usable part of the ingested energy, I (light in
case of autotrophs and food in case of heterotrophs) is assimilated, A while unusable part is ejected,
NU. A large portion of assimilated energy must be respired, R to provide existence energy to keep
the body functioning and repaired. A part of energy can be used for growth and reproduction, P,
while a part can be stored, S to accept further inputs of energy. Energy partitioning between P and R
is of vital importance to the individual and species.

Figure 3. A “universal” model of energy flow in a generalized ecosystem (based

on Odum 1963). I = input or ingested energy; NU = not utilized energy; A =
assimilated energy; P = production; R = respiration; B = biomass; G = growth; S =
stored energy; E = excreted energy

Primary producers convert the energy derived from sun into chemical energy in the form of food.
This creates two major kinds of energy pools, the living organic matter (plant biomass) and the non-
living organic matter (plant detritus). These pools form the base of two major pathways of energy
transfers (Figure 4). The biophagic or grazing pathway comprises the ingestion of living organic

Ecosystem Ecology
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Ecosystem Function
 matter by herbivores, while in the detritus or saprophagic pathway, detritus is consumed by the
detritivores or saprovores. Organisms following the biophagic pathway constitute biophagic
subsystem, and those that follow saprophagic pathway comprise the decomposition subsystem in an
ecosystem. All heterotrophic organisms depending on food produced by autotrophs (primary
producers) are called secondary producers and the organic matter synthesized by them constitutes
secondary production.

Figure 4. Energy pools and associated pathways of energy transfer

Food chain and food web

A large number of microscopic phytoplanktons and other macrophytes perform the primary job of
energy fixation. They, the primary producers are eaten by herbivores, which are further eaten by
carnivores or predators in different ecosystems. The transfer of energy fixed by plants through a
series of organisms eating one and being eaten by other is called food chain. A typical example of a
food chain can be traced in a grassland ecosystem as shown in Figure 5.

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 Grasses Grasshoppers Frogs-Snakes Vultures

Figure 5. Flow of energy in a generalized grazing food chain

However, the food relations are not so simple; rather they are complex. The trophic relationship
between organisms of any ecosystem is not always in a simple chain-like fashion but forms a
complicated network. This net-like trophic inter-relationship is called a food web. Since several
species may occur in each of the trophic levels, the organisms that feed on a common set of
organisms and are fed on by another common set of organisms are referred to as trophic species
(Briand and Cohen 1984). The analysis of food webs is important to understand ecosystem
dynamics in terms of limited euryphagy and euryphagous primary and secondary consumers (Figure
6). According to Paine (1980), three types of food webs can be depicted, namely, connectedness
webs- based on the concept of ‘who eats whom’, emphasizing the feeding relationships, energy flow
food webs- where energy flow through the food web is estimated and quantified by the flux of
energy between a resource and its consumer, and functional food webs- developed the idea of
functional food webs on the basis of impact of species on the structure of community. The food
webs have also been described as source webs- refer to one or more kinds of organisms that eat
them, their predators, and so on, sink webs- refer to one or more kinds of organisms, the organisms
they eat, their other prey, and so on (descending trophic levels), and community webs- represents a
group of species within a defined area of habitat.

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 Figure 6. Inter-linked network of food chains and food webs in an ecosystem

Ecological pyramids

There exists a relationship between the numbers, biomass, and energy contents of the primary
producers, first and second order consumers, and so on to the top carnivores in any ecosystem. Such
relationships represented in diagrammatic ways are referred to as ecological pyramids. Ecological
pyramids are of three categories, namely, of numbers, of biomass, and of energy or productivity.
Ecological pyramid of numbers and biomass may be upright pyramidal, inverted pyramidal or of
any other shape, but the pyramid of energy is always upright pyramidal or triangle-shaped.

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 Figure 7. Pyramid of numbers in a grassland ecosystem

Pyramid of numbers is the one in which individuals at each of the successive trophic levels are
counted per unit area and their numbers are plotted (Figure 7), pyramid of biomass, in which the
total biomass existing at each of the successive trophic levels is measured in terms of dry weight or
caloric value, per unit area and plotted (Figure 8 & 9), while pyramid of energy or productivity, in
which the energy flow per unit time at each of the successive trophic levels is measured and plotted
(Figure 10). The pyramid of energy flow is considered the one of fundamental significance as it
illustrates the actual functional relationships between the trophic levels. Being governed by the
second law of thermodynamics, energy flow declines from the producer level to successive trophic
levels. Consequently, the pyramid of energy necessarily undertakes a typical upright pyramidal
shape, reflecting a stepwise decrease in energy or productivity towards increasing trophic level.
However, the shape of the pyramid of energy is not affected by size or rate of metabolism of
organisms, while the other two pyramids are affected.

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 Figure 8. Pyramid of biomass in a grassland ecosystem

Figure 9. Pyramid of biomass for various ecosystems

Animals may have larger biomass per unit area than plants, but their production per unit time per
unit area would be much smaller than that of plants. An energy pyramid is a presentation of the
trophic levels in an ecosystem. Energy from the sun is transferred through the ecosystem by passing
through various trophic levels. Roughly 10% of the energy is transferred from one trophic level to
the next, thus preventing transfer to a large number of trophic levels (see Figure 10). There must be
higher amounts of biomass at the bottom of the pyramid to support the energy and biomass
requirements of the higher trophic levels (see Figure 11).

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 Figure 10. Pyramid of energy in a grassland ecosystem

Figure 11. Comparison of pyramid of energy, biomass and number in an old field in Michigan
(modified from Odum 1971)

Nutrient cycling
The two processes proceed concurrently in ecosystems, the unidirectional flow of energy and the
cycling of nutrient elements. Life sustenance is not only dependent on energy but also on the
availability of key elements important for different life processes. Importantly, flow of energy is
unidirectional and non-cyclic, whereas the movement of nutrient elements is cyclic. The conversion
of radiant energy into chemical energy during photosynthesis, various inorganic elements and
compounds are incorporated into the protoplasm of producers. The notable ones include the direct
components of photosynthetic reaction, carbon dioxide, water, and those that are critical for
protoplasmic synthesis viz. nitrogen, phosphorus, sulfur, magnesium (macronutrients), and other
essential nutrients needed in small quantities (micronutrients). In the process of consumption of

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 green plant, not only stored chemical energy in the form of carbohydrates, fats, and proteins is
transferred to the herbivores, but nutrients are also transferred. Similarly, both, energy and nutrients
are transferred from herbivores to carnivores and from the preceding trophic levels to the
decomposers. Thus, the flow of energy and nutrients goes hand-in-hand in an ecosystem in a
broadly similar pattern.
Although, there is a progressive decrease of energy in a food chain, the nutrients are not diminished.
Eventually, nutrient-containing protoplasm is subjected to decomposition that releases nutrients in
the environment and makes them available for reuse and recycling. The nature of the nutrient
cycling process is closely inter-twined with the interaction of physical, chemical, and biotic factors
in an ecosystem. Ecosystems, being open systems participate in various biogeochemical processes
through a system of inputs and outputs. Broadly, there are two inter-related nutrient budget systems
in an ecosystem, namely intra-system cycling of nutrients and extra-system nutrient transfers
(Figure 12). The intra-system cycling is concerned with the circulation of nutrients through the
biotic and abiotic compartments of an ecosystem. It involves the input and output along the
different trophic levels and the exchanges between reservoirs and sediments of an ecosystem.
However, the extra-system transfers pertain to the intake and output of the entire ecosystem in
relation to other ecosystems.
Productivity
The pyramid of energy introduces a very important factor, i.e. the rate. The rate of energy fixation
by green plants regulates the rate of production of organic material from simple inorganic
substances in a given area over a given period of time. This rate of energy fixation and consequent
increase in produced organic biomass is called primary productivity. Hence, the primary
productivity can be defined as the primary fixation of radiant energy by plants in the form of
organic substances and subsequent use of the fixed energy by herbivores, carnivores, and
detritivores. Broadly, total amount of energy that is fixed by plants in a unit area in a given period
of time is referred as gross primary productivity. A significant part of this energy is released during
plant respiration for use in metabolic activities. Hence, net primary productivity is smaller because
it is adjusted for energy losses required to support plant respiration and defined as the rate of
storage of energy (or organic matter) in plant tissues when loss due to respiratory utilization by the

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 Figure 12. A generalized model of intra-system nutrient cycling and extra-system nutrient transfer

plants is deducted out of the gross production rate. From producers, a portion of fixed energy gets
transferred to heterotrophs that ingest living matter, while the remaining is converted to detritus
pool to support those heterotrophs that feed on non-living organic matter. Within food webs, a
pyramid-shaped structure characterizes ecological productivity. Plants typically account for more
than 90% of the total productivity of the food web, herbivores most of the rest, and carnivores less
than 1%.
Various factors influence the rate and amount of energy fixation during photosynthesis viz. the
availability of basic chemical components of photosynthesis (moisture, nutrients, dissolved
substances, etc.), diurnal and seasonal changes in physical and biological factors, species diversity
in an ecosystem, etc. Because of differences in the availabilities of different factors, the world's
ecosystems differ greatly in the amount of productivity that they sustain. Sustainable ecosystem
relies on balance between the resource consumption and production. Unsustainable use of resources
feeds back on itself and the ecosystem loses productivity in the long term.

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 References
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ecological structure and functioning into designation of Special Areas of Conservation (SAC)
in the 0-12 nautical mile zone. School of Biological Sciences, University of Liverpool. (2006)
158pp.
Giller, P. S., Hillebrand, H., Berninger, U.-G. et al. 2004. Biodiversity effects on ecosystem
functioning: emerging issues and their experimental test in aquatic environments. Oikos 104:
423-436.
Muller, F. 1997. State-of-the-art in ecosystem theory. Ecol. Model. 100:135-161.
Myster, R.W. 2001. What is Ecosystem Structure?. Caribb. J. Sci. 37: 132-134.
Odum, E. P. 1953. Fundamentals of Ecology. Saunders, Philadelphia. First edition, 384 pp.
Odum, E.P., Barrett, G.W. 2005. Fundamentals of Ecology (5th ed.). Thomson, Brooks/Cole:
Australia.
Tansley, A. G. 1935. The use and abuse of vegetational concepts and terms. Ecology 16: 284-307.
Waide, R.B. 1999. The Relationship between Productivity and Species Richness. Annu. Rev. Ecol.
Syst. 30: 257-300.

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Ecosystem Function