Introduction

Organisms of different species can interact in many ways. They can compete, or they can be symbionts

longterm partners with a close association. Or, of course, they can do what we so often see in nature

programs: one of them can eat the otherchomp! That is, they can form one of the links in a food chain.

In ecology, a food chain is a series of organisms that eat one another so that energy and nutrients flow

from one to the next.

Autotrophs vs. heterotrophs

What basic strategies do organisms use to get food? Some organisms, called autotrophs, also known as

self-feeders, can make their own foodthat is, their own organic compoundsout of simple molecules

like carbon dioxide. There are two basic types of autotrophs:

Photoautotrophs, such as plants, use energy from sunlight to make organic compoundssugarsout of

carbon dioxide in photosynthesis. Other examples of photoautotrophs include algae and cyanobacteria.

Chemoautotrophs use energy from chemicals to build organic compounds out of carbon dioxide or

similar molecules. This is called chemosynthesis. For instance, there are hydrogen sulfide-oxidizing

chemoautotrophic bacteria found in undersea vent communities where no light can reach.

Autotrophs are the foundation of every ecosystem on the planet. That may sound dramatic, but it's no

exaggeration! Autotrophs form the base of food chains and food webs, and the energy they capture from

light or chemicals sustains all the other organisms in the community. When we're talking about their role

in food chains, we can call autotrophs producers.

Heterotrophs, also known as other-feeders, can't capture light or chemical energy to make their own food

out of carbon dioxide. Humans are heterotrophs. Instead, heterotrophs get organic molecules by eating

other organisms or their byproducts. Animals, fungi, and many bacteria are heterotrophs. When we talk

about heterotrophs' role in food chains, we can call them consumers. As we'll see shortly, there are many
 different kinds of consumers with different ecological roles, from plant-eating insects to meat-eating

animals to fungi that feed on debris and wastes.

Food chains

Now, we can take a look at how energy and nutrients move through a ecological community. Let's start

by considering just a few who-eats-who relationships by looking at a food chain.

A food chain is a linear sequence of organisms through which nutrients and energy pass as one organism

eats another. Let's look at the parts of a typical food chain, starting from the bottomthe producersand

moving upward.

At the base of the food chain lie the primary producers. The primary producers are autotrophs and are

most often photosynthetic organisms such as plants, algae, or cyanobacteria.

The organisms that eat the primary producers are called primary consumers. Primary consumers are

usually herbivores, plant-eaters, though they may be algae eaters or bacteria eaters.

The organisms that eat the primary consumers are called secondary consumers. Secondary consumers

are generally meat-eaterscarnivores.

The organisms that eat the secondary consumers are called tertiary consumers. These are carnivore-

eating carnivores, like eagles or big fish.

Some food chains have additional levels, such as quaternary consumerscarnivores that eat tertiary

consumers. Organisms at the very top of a food chain are called apex consumers.

We can see examples of these levels in the diagram below. The green algae are primary producers that get

eaten by mollusksthe primary consumers. The mollusks then become lunch for the slimy sculpin fish, a

secondary consumer, which is itself eaten by a larger fish, the Chinook salmona tertiary consumer.
 In this illustration, the bottom trophic level is green algae, which is the primary producer. The primary

consumers are mollusks, or snails. The secondary consumers are small fish called slimy sculpin. The

tertiary and apex consumer is Chinook salmon.

Image credit: Ecology of ecosystems: Figure 3 by OpenStax College, Biology, CC BY 4.0

Each of the categories above is called a trophic level, and it reflects how many transfers of energy and

nutrientshow many consumption stepsseparate an organism from the food chain's original energy

source, such as light. As well explore further below, assigning organisms to trophic levels isn't always

clear-cut. For instance, humans are omnivores that can eat both plants and animals.

Decomposers
One other group of consumers deserves mention, although it does not always appear in drawings of food

chains. This group consists of decomposers, organisms that break down dead organic material and

wastes.

Decomposers are sometimes considered their own trophic level. As a group, they eat dead matter and

waste products that come from organisms at various other trophic levels; for instance, they would happily

consume decaying plant matter, the body of a half-eaten squirrel, or the remains of a deceased eagle. In a

sense, the decomposer level runs parallel to the standard hierarchy of primary, secondary, and tertiary

consumers.

Fungi and bacteria are the key decomposers in many ecosystems; they use the chemical energy in dead

matter and wastes to fuel their metabolic processes. Other decomposers are detritivoresdetritus eaters

or debris eaters. These are usually multicellular animals such as earthworms, crabs, slugs, or vultures.

They not only feed on dead organic matter but often fragment it as well, making it more available for

bacterial or fungal decomposers.

Examples of decomposers: left, fungi growing on a log; right, an earthworm.

Image credit: left, Decomposers by Courtney Celley/USFWS, CC BY 2.0; right, Earthworm by Luis
Miguel Bugallo Snchez, CC BY-SA 3.0

Decomposers as a group play a critical role in keeping ecosystems healthy. When they break down dead

material and wastes, they release nutrients that can be recycled and used as building blocks by primary

producers.
Food webs

Food chains give us a clear-cut picture of who eats whom. However, some problems come up when we

try and use them to describe whole ecological communities.

For instance, an organism can sometimes eat multiple types of prey or be eaten by multiple predators,

including ones at different trophic levels. This is what happens when you eat a hamburger patty! The cow

is a primary consumer, and the lettuce leaf on the patty is a primary producer.

To represent these relationships more accurately, we can use a food web, a graph that shows all the

trophiceating-relatedinteractions between various species in an ecosystem. The diagram below

shows an example of a food web from Lake Ontario. Primary producers are marked in green, primary

consumers in orange, secondary consumers in blue, and tertiary consumers in purple.

The bottom level of the illustration shows primary producers, which include diatoms, green algae, blue-

green algae, flagellates, and rotifers. The next level includes the primary consumers that eat primary

producers. These include calanoids, waterfleas, cyclopoids, rotifers and amphipods. The shrimp also eat

primary producers. Primary consumers are in turn eaten by secondary consumers, which are typically

small fish. The small fish are eaten by larger fish, the tertiary consumers. The yellow perch, a secondary

consumer, eats small fish within its own trophic level. All fish are eaten by the sea lamprey. Thus, the

food web is complex with interwoven layers.

Image credit: Ecology of ecosystems: Figure 5 by OpenStax College, Biology, CC BY 4.0; original work
by NOAA, GLERL

In food webs, arrows point from an organism that is eaten to the organism that eats it. As the food web

above shows, some species can eat organisms from more than one trophic level. For example, opossum

shrimp eat both primary producers and primary consumers.

Applications of Food Webs

Food webs are constructed to describe species interactions (direct relationships).
The fundamental purpose of food webs is to describe feeding relationship among species in a community.
Food webs can be constructed to describe the species interactions. All species in the food webs can be
distinguished into basal species (autotrophs, such as plants), intermediate species (herbivores and
intermediate level carnivores, such as grasshopper and scorpion) or top predators (high level carnivores
such as fox) (Figure 1).

These feeding groups are referred as trophic levels. Basal species occupy the lowest trophic level as
primary producer. They convert inorganic chemical and use solar energy to generate chemical energy.
The second trophic level consists of herbivores. These are first consumers. The remaining trophic levels
include carnivores that consume animals at trophic levels below them. The second consumers (trophic
level 3) in the desert food web include birds and scorpions, and tertiary consumers making up the fourth
trophic level include bird predators and foxes. Grouping all species into different functional groups or
tropic levels helps us simplify and understand the relationships among these species.

Food webs can be used to illustrate indirect interactions among species.

Indirect interaction occurs when two species do not interact with each other directly, but influenced by a
third species. Species can influence one another in many different ways. One example is the keystone
predation are demonstrated by Robert Paine in an experiment conducted in the rocky intertidal zone (Cain
et al. 2008; Smith & Smith 2009; Molles 2010). This study showed that predation can influence the
competition among species in a food web. The intertidal zone is home to a variety of mussels, barnacles,
limpets, and chitons (Paine 1969). All these invertebrate herbivores are preyed upon by the predator
starfish Pisaster (Figure 3). Starfish was relatively uncommon in the intertidal zone, and considered less
important in the community. When Paine manually removed the starfish from experimental plots while
leaving other areas undisturbed as control plots, he found that the number of prey species in the
experimental plots dropped from 15 at the beginning of the experiment to 8 (a loss of 7 species) two years
after the starfish removal while the total of prey species remained the same in the control plots. He
reasoned that in the absence of the predator starfish, several of the mussel and barnacle species (that were
superior competitors) excluded the other species and reduced overall diversity in the community (Smith &
Smith 2009). Predation by starfish reduced the abundance of mussel and opened up space for other
species to colonize and persist. This type of indirect interaction is called keystone predation.
Figure 3
(a) The rocky intertidal zone of the Pacific Northwest coast is inhabited by a variety of species including
starfish, barnacles, limpets, chitons, and mussels. (b) A food web of this community shows that the
starfish preys on a variety of invertebrate species. Removal of starfish from this community reduced the
diversity of prey species due to increased competition.
Another interesting study demonstrated indirect interactions among species in both aquatic and terrestrial
ecosystems (Figure 4). In a study conducted near Gainesville, Florida, Knight and her colleagues (2009)
investigated the effects of fish in ponds on plant seeds production. They measured and compared
abundances of both larval and adult dragonfly in and around four ponds that had been stocked with fish
and four ponds that lacked fish (Knight et al. 2009). They found that ponds with fish produce fewer larval
and adult dragonflies than ponds without fish, as fish prey on larval dragonflies. As dragonfly population
decreases, the populations of their prey, including bees, flies, and butterflies, decrease. These prey species
are pollinators of the plants. Therefore, flowers in the vicinity of ponds without fish receive fewer
pollinator visits than flowers close to ponds stocked with fish. Since the production of seeds is pollen-
limited, fewer pollinator visits result in lower seeds production. This study demonstrates, via a complex
trophic cascade, that adding fish to a pond improves the reproductive success of a plant on land (Ricklefs
2008).
Figure 4: An interaction food web shows that fish have indirect effects on the populations of several
species in and around ponds.
The solid arrows represent direct effects, and the dashed arrows indirect effects; the nature of the effect is
indicated by + or -. Fish have indirect effects, through a trophic cascade, on several terrestrial species:
dragonfly adults (-), pollinators (+), and plants (+)

2012 Nature Education All rights reserved.

Food webs can be used to study bottom-up or top-down control of community structure.
Food webs illustrate energy flow from primary producers to primary consumers (herbivores), and from
primary consumers to secondary consumers (carnivores). The structure of food webs suggests that
productivity and abundance of populations at any given trophic level are controlled by the productivity
and abundance of populations in the trophic level below them (Smith & Smith 2009). This phenomenon is
call bottom-up control. Correlations in abundance or productivity between consumers and their resources
are considered as evidence for bottom-up control. For example, plant population densities control the
abundance of herbivore populations which in turn control the densities of the carnivore populations. Thus,
the biomass of herbivores usually increases with primary productivity in terrestrial ecosystems.
Top-down control occurs when the population density of a consumer can control that of its resource, for
example, predator populations can control the abundance of prey species (Power 1992). Under top-down
control, the abundance or biomass of lower trophic levels depends on effects from consumers at higher
trophic levels. A trophic cascade is a type of top-down interaction that describes the indirect effects of
predators. In a trophic cascade, predators induce effects that cascade down the food chain and affect
biomass of organisms at least two links away (Ricklefs 2008). Nelson Hairston, Frederick Smith and
Larry Slobodkin first introduced the concept of top-down control with the frequently quoted "the world is
green" proposition (Power 1992; Smith & Smith 2009). They proposed that the world is green because
carnivores depress herbivores and keep herbivore populations in check. Otherwise, herbivores would
consume most of the vegetation. Indeed, a bird exclusion study demonstrated that there were significantly
more insects and leaf damage in plots without birds compared to the control (Marquis & Whelan 1994).
Food webs can be used to reveal different patterns of energy transfer in terrestrial and aquatic
ecosystems.
Patterns of energy flow through different ecosystems may differ markedly in terrestrial and aquatic
ecosystems (Shurin et al. 2006). Food webs (i.e., energy flow webs) can be used to reveal these
differences. In a review paper, Shurin et al. (2006) provided evidence for systematic difference in energy
flow and biomass partitioning between producers and herbivores, detritus and decomposers, and higher
trophic levels in food webs. A dataset synthesized by Cebrian and colleagues on the fate of carbon fixed
by primary productivity across different ecosystems was used to show different patterns in food chains
between terrestrial and aquatic ecosystems (Figure 5). On average, the turnover rate of phytoplankton is
10 to 1000 times faster than that of grasslands and forests, thus, less carbon is stored in the living
autotroph biomass pool, and producer biomass is consumed by aquatic herbivores at 4 times the terrestrial
rate (Cebrian 1999, 2004; Shurin et al. 2006). Herbivores in terrestrial ecosystems are less abundant but
decomposers are much more abundant than in phytoplankton dominated aquatic ecosystems. In most
terrestrial ecosystems with high standing biomass and relatively low harvest of primary production by
herbivores, the detrital food chain is dominant (Smith & Smith 2009). In deep-water aquatic ecosystems,
with their low standing biomass, rapid turnover of organisms, and high rate of harvest, the grazing food
chain may be dominant.

Figure 5: Differences in pathways of carbon flow and pools between aquatic and terrestrial
ecosystems.
The thickness of the arrows (flows) and the area of the boxes (pools) correspond to the magnitude. The
size of the pools are scaled as log units since the differences cover four orders of magnitude. The Cs
indicate consumption terms (i.e. CH is consumption by herbivores). Ovals and arrows in grey indicate
unknown quantities.
As a diagram tool, food web has been approved to be effective in illustrating species interactions and
testing research hypotheses. It will continue to be very helpful for us to understand the associations of
species richness/diversity with food web complexity, ecosystem productivity, and stability.
Energy transfer efficiency limits food chain lengths

Energy is transferred between trophic levels when one organism eats another and gets the energy-rich

molecules from its prey's body. However, these transfers are inefficient, and this inefficiency limits the

length of food chains.

When energy enters a trophic level, some of it is stored as biomass, as part of organisms' bodies. This is

the energy that's available to the next trophic level since only energy storied as biomass can get eaten. As

a rule of thumb, only about 10% of the energy that's stored as biomass in one trophic levelper unit

timeends up stored as biomass in the next trophic levelper the same unit time. This 10% rule of

energy transfer is a good thing to commit to memory.

Trophic pyramid illustrating the 10% energy transfer rule.

Light energy is captured by primary producers.

Amount of energy stored as biomass:

Primary producers20,000 kcal per meter squared per year

Primary consumers2,000 kcal per meter squared per year

 Secondary consumers200 kcal per meter squared per year

Tertiary consumers20 kcal per meter squared per year

Tertiary consumers2 kcal per meter squared per year

At each level, energy is lost directly as heat or in the form of waste and dead matter that go to the

decomposers. Eventually, the decomposers metabolize the waste and dead matter, releasing their energy

as heat also.

Image credit: modified from Ecological pyramid by CK-12 Foundation, CC BY-NC 3.0

Why does so much energy exit the food web between one trophic level and the next? Here are a few of

the main reasons for inefficient energy transfer^{1,2}1,2start superscript, 1, comma, 2, end superscript:

In each trophic level, a significant amount of energy is dissipated as heat as organisms carry out cellular

respiration and go about their daily lives.

Some of the organic molecules an organism eats cannot be digested and leave the body as feces, poop,

rather than being used.

Not all of the individual organisms in a trophic will level get eaten by organisms in the next level up.

Some instead die without being eaten.

The feces and uneaten, dead organisms become food for decomposers, who metabolize them and convert

their energy to heat through cellular respiration. So, none of the energy actually disappearsit all winds

up as heat in the end.