Ecological Homeostasis

Ecological Homeostasis

Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Ecological Homeostasis
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.

All rights reserved. No part of this publication may be reproduced, stored

in a retrieval system, or transmitted in any form or by any means
electronic, mechanical, photocopy, recording, or any otherexcept for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.

First published in 2016 by

Momentum Press, LLC
222 East 46th Street, New York, NY 10017
www.momentumpress.net

ISBN-13: 978-1-60650-953-1 (print)

ISBN-13: 978-1-60650-954-8 (e-book)

Momentum Press Biology Collection

Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

Chennai, India

10 9 8 7 6 5 4 3 2 1

Printed in the United States of America

 Abstract
Individual organisms contribute to nutrient cycling in ecological sys-
tems, which is shown to be a mechanism of homeostasis at that level. The
phosphorus and nitrogen cycles are used to illustrate effects of changes
in populations or communities on the cycling of these nutrients. Major
disturbances such as deforestation and global climate change disrupt
nutrient cycles and ecological system homeostasis. Data are examined to
determine effects of deforestation on nutrient cycling. Increasing atmo-
spheric carbon dioxide and global climate change are disrupting ecologi-
cal systems homeostasis, and several studies are used to show how this is
happening, including changes in primary production, temperature and
precipitation patterns. This book also discusses the role of individual spe-
cies in filtering contaminants and pollutants from ecological systems.

Keywords
nitrogen cycle, assimilation, decomposition, nitrogen fixation, ecological
system, nutrient dynamics, biomass, nutrient regeneration, pollutants,
contaminants, hyperaccumulators, tragedy of the commons, greenhouse
gas, carbon sink, carbon source, net primary production, gross primary
production, respiration, ecological mismatch, positive feedback
 Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Nutrient Cycling Is a Mechanism of Homeostasis
for Ecological Systems........................................................1
Ethical, Legal, Social Implications: The Gulf
of Mexico Dead Zone Is Related to Increased
Nutrient Input..............................................................14
Chapter 2 Ecological Systems Can Filter Wastes Like Individual
Organisms........................................................................19
Ethical, Legal, Social Implications: Pollution Is a
Tragedy of the Commons.............................................24
Chapter 3 Increasing Atmospheric Carbon Dioxide Can Disrupt
Ecological Systems...........................................................27
Ethical, Legal, Social Implications: There Is a
Difference Between Weather and Climate.....................40
Conclusion............................................................................................43
Glossary................................................................................................45
Index....................................................................................................47
 Preface
This book about ecological system homeostasis is part of a thirty book
series that collectively surveys all of the major themes in biology. Rather
than just present information as a collection of facts, the reader is treated
more like a scientist, which means the data behind the major themes are
presented. Reading any of the thirty books by Paradise and Campbell
provides readers with biological context and comprehensive perspective
so that readers can learn important information from a single book with
the potential to see how the major themes span all size scales: molecular,
cellular, organismal, population and ecologic systems. The major themes
of biology encapsulate the entire discipline: information, evolution, cells,
homeostasis and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, readers will learn about several examples of ecological system
homeostasis and some of the supporting evidence behind our understand-
ing. The historic and more recent experiments and data will be explored.
Instead of believing or simply accepting information, readers of this book
will learn about the science behind ecological system homeostasis the way
professional scientists dowith experimentation and data analysis. In
short, data are put back into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this
content can go to www.bio.davidson.edu/icb where they will find
pedagogically-designed and interactive Integrating Concepts in Biology
for introductory biology college courses or a high school AP Biology
course.
 Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with co-author AMC. Davids gift allowed us to hire talented artists (Tom
Webster and his staff at Lineworks, Inc.) and copyeditor Laura Loveall.
Thanks go to Kristen Mandava of Mandava Editorial Services for project
management and guidance. In particular, we are indebted to Katie Noble
and Melissa Hayban for their many hours and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to
administrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
Thanks to my wife Amy Brooks for her constant support during the
development of this textbook, and my daughter Evelyn for her endless
energy. Thanks to Malcolm Campbell for his steadfast resolve and opti-
mism. Without him, this book would not exist. Thanks to collaborator
Laurie Heyer for taking my sometimes half-baked math ideas and turning
them into powerful and elegant Bio-Math Explorations. I learned a lot
from both of them. While the math is largely absent from this book, our
collaboration with her made this a better book. Nancy Stamp at Bing-
hamton University, and Bill Dunson and Richard Cyr at The Pennsyl-
vania State University influenced me greatly in how I think as a scientist
and approach my teaching. Finally, I thank my students in Integrated
Concepts in Biology II, who enthusiastically participated in our experi-
ment to redesign introductory biology, starting with the text and ending
with a new approach to teaching biology.
 Introduction
For most people, it is difficult to think of an entire ecological system,
comprised of a multitude of species, maintaining homeostasis. But if one
thinks of a forest, one that they had visited year after year, they may realize
that it has not changed much over the years. In this book, negative feed-
back mechanisms that operate at the level of the ecological system will be
examined. A possible exception to maintenance of homeostasis is when
an ecological system is subjected to a large scale disturbance, which has
disrupted homeostasis. Think about coastal ecological systems buffeted by
hurricanes. It may take much time and energy for processes acting in eco-
logical systems to return the system to its original state. Alternatively, as
with populations, changes in the environment can trigger changes in the
stable state that had been maintained by negative feedback mechanisms.
The larger size of these systems and their environment influence how they
address physical and chemical challenges. Emergent properties play a key
role in maintaining homeostasis in ecological systems; the mechanisms
may be caused by individual species maintaining their populations or
individuals gathering energy and nutrients. Because life requires energy
and nutrients, a lot can be learned about ecological system homeostasis by
examining nutrient cycling.
 CHAPTER 1

Nutrient Cycling Is a
Mechanism of Homeostasis
for Ecological Systems

In one step of the nitrogen cycle, molecular nitrogen moves from the
atmosphere to Rhizobium bacteria, which convert it to ammonia and
then incorporate it into their amino acids and provide it to legumes.
Assimilation by organisms, which results in production of organic forms
of nitrogen (proteins and DNA), and decomposition, which reverses
assimilation, further move nitrogen through the cycle. Nitrogen fixation,
assimilation, and decomposition are the mechanisms that cycle nitrogen
through ecological systems.
More generally, nutrient cycles describe the mechanisms by which
nutrients are altered and transferred between parts of ecological systems.
These parts are broadly considered compartments in ecological systems.
A compartment is a part or space into which an ecological system is sub-
divided. The biotic compartment consists of all the biological organisms
in the ecological system, although biologists are often interested in the
nutrients that flow between individual species or from primary producers
to herbivores. Other compartments include the soil, water, and air. Move-
ment often includes transformation of the element, from one form to
another. For instance, the mechanism of photosynthesis simultaneously
moves carbon from the atmosphere to the primary producers and changes
the form of carbon from carbon dioxide to a carbohydrate.
Some studies have examined the role of individual species in nutri-
ent cycles. Michael Vanni and his colleagues studied nutrient dynamics
in Tuesday Lake in Michigan (Vanni et al., 1997). Nutrient dynamics
are the changes in nutrient concentrations in compartments over time.
2 ECOLOGICAL HOMEOSTASIS

The scientists were interested in the effects of fish on nutrient dynamics,

and they were specifically interested in how fish that eat zooplankton
(tiny, free-floating aquatic animals) affect the cycling of nitrogen (N) and
phosphorus (P), the location of the nutrients, and what mechanisms are
involved in movement and transformation of nutrients. Only the phos-
phorus cycle will be examined here, although Vanni and colleagues also
studied the nitrogen cycle.
The scientists conducted enclosure experiments to quantify the effects
of fish on nutrient dynamics. An enclosure is a device, fence, or container
that maintains certain species within an ecological system. Enclosures
were suspended from the surface of the lake and were made of cylindrical
plastic tubes, which the scientists closed off from the lake at the bottom
and left open at the top. These plastic bags were 1 meter in diameter and
3 meters deep with a volume of over 2,000 liters. The scientists filled all
enclosures with water filtered to remove zooplankton. Known amounts
of zooplankton were added to each enclosure. The experiment was per-
formed twice. In the first summer, the scientists used four treatments: no,
low, medium, and high fish, with each treatment replicated three times.
In the second summer, three treatments were used (no, low, and high fish)
with four replicates per treatment. The estimated mass of minnows per
hectare of lake surface area for Tuesday Lake was about 50. The average
mass of minnows in the no fish treatment was 0, of course, for low fish it
was 32.5 kg/ha (year 1, with 2 fish added) and 46.7 kg/ha (year 2, with
3 fish added), for medium fish it was 66.2 (year 1, with 4 fish added), and
for high fish it was 113.9 (year 1, with 7 fish added) and 97.7 (year 2,
with 6 fish added). The scientists captured minnows that eat zooplankton,
placed them in buckets, measured them, and added them to enclosures
within 1 hour of being captured.
Enclosures were sampled before, during, and at the end of the tri-
als, depending upon the variable being measured. Vanni and his col-
leagues quantified the concentration of phosphorus in the water column,
including total and particulate fractions. The water column is open
waterbetween the surface and the sediment. Particulates are living or
nonliving material suspended in the water column. Water column phos-
phorus represents the total phosphorus suspended in the water column,
including the amount dissolved in water, and the amount incorporated
 Nutrient Cycling Is a Mechanism of Homeostasis 3

in zooplankton, phytoplankton, bacteria, and detritus but excluding the

fish. The scientists also quantified particulate phosphorus separately from
total phosphorus. Water samples were filtered and dried. In the second
year, the samples were additionally filtered to separate out a smaller frac-
tion of particulate matter, called seston. Seston is minute living and
nonliving matter (<63 mm) suspended in the water column. All particu-
late matter was assayed for phosphorus.
The biologists sampled zooplankton in the water column twice per
week. They sampled at five depths in each enclosure, pooled all zooplank-
ton, and then analyzed the sample for species composition, mass, and
phosphorus concentration. Vanni and his colleagues estimated density,
and then they estimated mass of zooplankton by measuring the size of
many individuals of each species and converting length to wet mass using
known relationships; biomass was assumed to be 5% of wet mass. Indi-
vidual biomass was then converted to population biomass, based on the
estimated population density (number/liter) times the mean biomass.
Vanni and his colleagues sampled phytoplankton twice per week from
each enclosure and from outside the enclosures.
Strips of plastic hung on the inside walls of the enclosure were used
to collect and sample organisms growing on the walls. The strips were
removed at the end of each years experiment; the algae and bacteria
growing on the plastic were removed, filtered, dried, and analyzed for
phosphorus (Figure 1). Material that settled out of the water column was
collected in traps placed at the bottom of enclosures at regular intervals,
filtered, dried, and analyzed as other samples (see Figure 1).
The biologists collected the minnows from the enclosures at the end
of the experiment and measured their length and mass. The dried fish
remains were ground into a powder, and samples were analyzed for phos-
phorus concentrations. Concentrations per unit biomass were multiplied
by fish biomass per enclosure to estimate the total amount of phosphorus
present in fish. The scientists estimated phosphorus concentration in fish
at the beginning of the experiment using a group of ten minnows not used
in enclosures but collected at the beginning of the experiment. They then
used those concentrations to estimate the initial concentrations in enclo-
sure fish based on the mass of those fish, and from that, the change in fish
phosphorus from beginning to end of the experiment (see Figure 1).
4 ECOLOGICAL HOMEOSTASIS
movement of P in and out of water column (mmoles/L 1SE)
100

100

200

= loss from WC to wall

= loss from WC to sediment
= loss from fish to WC
300 = calculated change in WC P
= actual change in WC P

no fish low fish medium fish high fish

Figure 1 Movement of phosphorus relative to the water column (WC)

for the year 1 enclosure experiment. Means are mmoles of P/liter over
the entire experiment (44 days). Error bars equal 1 standard error
(SE). Negative values indicate phosphorus left the water column
and accumulated in indicated compartment. Fish lost phosphorus,
which accumulated in the water column. Calculated change in water
column P is the sum of first three bars in each group.
Source: Data from Vanni et al., 1997.

Vanni and his colleagues found that fish increased phosphorus in the
water column. Part of that was due to fish losing mass in enclosures.
However, the researchers found fish lost less relative to the amount of
body mass lost, indicating that fish can conserve phosphorus under the
conditions that caused the loss of body mass. This may be important to
maintaining their individual homeostasis.
The proportion of total phosphorus present as particulates also tended
to be higher in fish presence. At the end of the second year experiment,
zooplankton constituted a smaller part of the particulate phospho-
rus in the low and high fish treatments than in the no fish treatment.
The smaller particles separated from zooplankton constituted a greater
 Nutrient Cycling Is a Mechanism of Homeostasis 5

fraction of particulate phosphorus when fish were present than when

fish were absent. This may be caused by fish reducing total zooplankton
numbers and preferentially consuming larger organisms in the plankton,
leaving the smaller zooplankton to dominate. The researchers concluded
that fish consuming zooplankton increase total phosphorus in both the
water column and particulate matter. In addition, phosphorus lost from
the water to assimilation by organisms growing on walls and through
sedimentation both increased with increasing fish biomass.
The loss of fish mass is an artifact of the experiment, because the min-
now studied typically eats prey near the shore, and they were prevented
from eating such prey while in enclosures. However, one can still learn
about the phosphorus cycle by considering phosphorus dynamics in
enclosures with different amounts of fish. Fish excrete phosphorus and
contribute it to the water column. The transfer of phosphorus from the
zooplankton to the water column, through the fish, supplies phosphorus
to phytoplankton. Under normal circumstances, fish that feed near shore
would transfer phosphorus from the animals near the shore to the phyto-
plankton in the water column, becoming a net source, rather than a sink,
of phosphorus.
The decline in total phosphorus over the course of the experiment was
similar in enclosures and the lake as a whole. If total phosphorus in the
water column declines during a growing season, there must be a mecha-
nism to restore it; otherwise it would continue to decrease. Particulate
nutrient concentrations were generally lower in the lake than in enclo-
sures, although differences between the lake and the no fish treatment
were generally small. The enclosure results may represent what happens
in a freshwater lake on a small spatial and temporal scale, especially in
enclosures where fish are present. That is, phosphorus cycled faster when
fish were present. Certainly, excretions from fish will replenish phospho-
rus and other nutrients, as Vanni and his colleagues showed. This is likely
to be a significant pathway in the phosphorus cycle (Figure 2). Through
similar experiments, scientists have come to better understand the phos-
phorus cycle. Accounting for inputs and outputs through the use of phos-
phorus budgets has been an important component of such experiments.
Vanni and his colleagues constructed such a budget for the first
year experiment, attempting to account for all phosphorus in each
6 ECOLOGICAL HOMEOSTASIS

= sedimentation
= assimilation rock
= erosion
= regeneration
= excretion
= decomposition

fish
carnivores

zooplankton
herbivores

algae
plants
water

terrestrial

sediment soil

Figure 2 Simplified schematic of the phosphorus cycle. Inorganic

forms are found in rock, sediment, and soil; and organic forms of
phosphorus are found in all biota compartments. The arrows for
decomposition represent dead organisms or parts of organisms that
decompose in the sediments and soils. The decomposer biota is not
shown.
Source: by C. Paradise.

compartment of the enclosure (in the water column, in organisms grow-

ing on the walls, in bottom sediment, and through fish biomass loss or
gain) and to determine how fish affected the fate of phosphorus and
the loss of phosphorus from the water column. The phosphorus budget
requires knowledge of how phosphorus moves through the environment
in the phosphorus cycle. Phosphorus leaves the water column through
the processes of sedimentation and uptake, or assimilation, by organisms
 Nutrient Cycling Is a Mechanism of Homeostasis 7

growing on the walls. It enters the water column through fish excretion.
These are three steps involved in the phosphorus cycle illustrated by Vanni
and his colleagues experiment.
Phosphorus budgets were found to be closely balanced in enclosures
with fish. The sum of losses from wall growth and sedimentation plus
gains from fish yielded a predicted value that was very close to the actual
change in water column phosphorus (see Figure 1). However, the budget
for fishless enclosures was highly imbalanced, perhaps due to an underes-
timation of phosphorus sinks. The predicted water column phosphorus
decline based on sedimentation and wall growth is only 57% of the actual
water column decline.
The researchers concluded that large zooplankton, more prevalent
in the no fish treatment, could more easily escape the sampling devices.
If more large zooplankton had been caught, the actual change in water
column phosphorus would have been less. Additionally, the amount of
phosphorus that accumulated on walls may have been underestimated in
no-fish enclosures. The scientists observed loosely attached algal growth
on walls of those enclosuresmuch more so than on fish enclosures.
These growths may have sloughed off as the plastic strips used to assess
wall growth were sampled. Despite these potential biases, results from
enclosure studies reveal it is possible to account for nutrients in such
experiments and can help scientists understand the role of individual spe-
cies in maintaining homeostasis in ecological systems.
The results of this experiment, as well as many others, indicate that fish
can have significant effects on the phosphorus cycle in lakes. In addition
to the steps described earlier (assimilation, sedimentation, and excretion),
there are other important steps in the phosphorus cycle (see Figure2).
The ultimate source of phosphorus is rock, which erodes and adds new
phosphorus to water and soil, replacing that which is lost due to sedimen-
tation from the water column and leaching from the soil. Decomposition
is another major process, where phosphorus is removed from dead organ-
isms and assimilated by decomposers. Regeneration occurs when phos-
phorus lost to the sediment is released by decomposers in inorganic form
and reenters the water column. Nutrient regeneration is the recycling and
release of nutrients from organic matter by decomposer organisms. This
brief overview of the phosphorus cycle shows the inputs and outputs of
8 ECOLOGICAL HOMEOSTASIS

a nutrient in multiple compartments. The balance of inputs and out-

puts in a compartment relate to the homeostasis of an ecological system;
although inputs and outputs do not always balance at any one time, they
often balance when averaged over time.
The phosphorus cycle does not occur in isolation in one ecological
system. Nutrients can, and do, move from one compartment to another,
and often those compartments are in different ecological systems. Move-
ment of nutrients from one ecological system to another partly accounts
for imbalances within a system. For instance, water running across the
land surface brings nutrients from the soil and vegetation to rivers. If
scientists only examined the land, they would find that nutrients are
being lost from the system. Scientists must conduct experiments to
determine how nutrients are then regenerated in such systems. Nitro-
gen fixation, the conversion of molecular nitrogen to ammonium and
nitrate by organisms, might be a major input in such situations. Nitro-
gen fixation in the watershed must balance losses in order to still main-
tain homeostasis.
If not balanced by inputs, long-term losses could disrupt homeostasis
of ecological systems. Large-scale natural or human-caused disturbances
could also disrupt homeostasis of nutrients. Herbert Bormann, Gene
Likens, and their colleagues tested the effect of complete deforestation
on nutrient cycling within an ecological system and nutrient export and
erosion of particulate matter from a forest to a stream (Bormann et al.,
1974). One of their objectives was to determine whether a previously
undisturbed forest had homeostatic capacity to hold nutrients when
nutrient assimilation by plants was destroyed. They studied a small water-
shed in the White Mountains of New Hampshire.
The scientists had determined rates of nutrients cycling in a treated
and a reference watershed, which allowed for comparison. During one
fall, the hardwood forest of the treatment watershed was completely
destroyed. The scientists cut down all of the trees, saplings, and shrubs in
the 15.6 hectare watershed. The reference watershed was 13.2 hectares in
area. All of the cut material in the treatment watershed was left in place
so that it could decompose on the ground. While applying the treatment
to the watershed, Bormann, Likens, and their colleagues were careful to
minimize soil erosion by preventing disturbance of the soil surface. The
 Nutrient Cycling Is a Mechanism of Homeostasis 9

400

annual particulate matter export (kg dry mass per ha) = undisturbed-organic
= undisturbed-inorganic
= undisturbed-total
= clearcut-organic
300 = clearcut-inorganic
= clearcut-total

200

100

0
0 1 2 3 4 5
year from time of clearcut

Figure 3 Annual particulate matter output in kilograms of dry mass

of organic and inorganic materials per hectare of watershed for
5years post-clearcut. Undisturbed refers to the undisturbed forested
watershed. ha, Hectare.
Source: Data from Bormann et al., 1974, Table 3.

scientists further prevented regrowth of vegetation the following 3.5 years

by spraying the entire watershed with an herbicide.
Bormann, Likens, and their colleagues set up a channel through
which all water in the stream that drained the watershed flowed. This
allowed them to collect water samples as well as accurately measure
discharge. Discharge is the volume of water in a river flowing past a point
during a specific time interval. The scientists measured particulate mat-
ter output from the stream that drained each watershed for 5 years after
the treatment watershed was clearcut. They determined the total annual
particulate matter export from data they collected on water discharge and
mass of collected particulate matter in the water (Figure 3).
The scientists also determined the concentrations of various nutri-
ents in the particulate matter and concentrations dissolved in the water
(Figure 4). These outputs constituted the total chemical output from
the ecological system via stream drainage. Mean gross losses of chemical
10 ECOLOGICAL HOMEOSTASIS

120
= undisturbed-OPM
= undisturbed-IPM
= undisturbed-DC
100 = clearcut-OPM
= clearcut-IPM
mean annual export (kg per ha)

= clearcut-DC
80

60

40

20

0
A Ca N

30

25
mean annual export (kg per ha)

20

15

10

0
Fe Mg P K Na
B elemental nutrient

Figure 4 Mean annual export in kilograms per hectare (ha) of

watershed of various elements in organic (OPM) and inorganic
particulate matter (IPM) and net dissolved concentration
(DCis output in water minus input in precipitation) for 4 years
after clearcutting the treatment watershed.
Source: Data from Bormann et al., 1974, Table 4.
 Nutrient Cycling Is a Mechanism of Homeostasis 11

elements in particulate matter were obtained by multiplying data from

Figure 3 by the percentage of various elements in organic debris and inor-
ganic materials. Organic debris was made up of twig and branch debris,
leaves, bark, and fruits of various sizes and fine and very fine material,
mostly undistinguishable as to origin. Inorganic material consisted of
sand, silt, clay, and materials of geologic origin. Concentrations of dis-
solved substances were measured in water samples collected at the channel.
As the scientists determined, water running over the land surface
can carry loose particulate matter and erode soil. This runoff ends up
in streams, carrying with it the particulate matter from the watershed.
Surface runoff is a major component of the hydrologic cycle, and it
may increase stream discharge intermittently or seasonally after periods
of heavy rainfall or snowmelt. The hydrologic cycle is the circulation of
water throughout ecological systems and the atmosphere through evapo-
ration, transpiration, and precipitation. The increased water creates more
turbulence, which causes suspension of particulate matter. Bormann, Lik-
ens, and their colleagues concluded that brief, intense storms and the run-
off associated with those storms is a major factor in removing particulate
matter from both undisturbed and clearcut watersheds.
In undisturbed, forested watersheds, there is variation in particulate
matter export over time. The forested watershed had an output of about
25 kg hectare21 of particulate matter, averaged over the 6 years of the
study. Output ranged from 7 to 49 kg hectare21 year21, and the
scientists concluded that variation was caused by the occurrence and tim-
ing of high river flows during particular years. This was tied to variable
precipitation in the watershed. For the forested ecosystem, storm periods
had a weighted average of 11.2 kg of particulate matter per 106 liters of
runoff, compared to an overall average of 2.2 kg per 106 liters.
Clearcutting the forest had a dramatic effect on particulate matter
export relative to undisturbed, forested watersheds. Total particulate mat-
ter export from the clearcut watershed averaged 156 kg hectare21
year21, which was six times higher than the output of the undisturbed
watershed. Export was highly variable, ranging from 16 to 380 kg
hectare21 year21, and increased each year following deforestation up to
year 4. In addition to increasing over time, the clearcut watershed was also
more susceptible to intense runoff. Storm periods produced a weighted
12 ECOLOGICAL HOMEOSTASIS

average of 67 kg of particulate matter per 106 liters of runoff as opposed

to an overall average output of 14.6 kg per 106 liters. There was a decrease
in export in year 5, which was back to levels seen in year 3. The researchers
speculated that this was probably due to previous erosion of much of the
available particulate matter, and regrowth of vegetation that occurred dur-
ing the final year when herbicide applications were no longer occurring.
Overall, for both watersheds, much of the total particulate matter,
particularly the very large particles, was exported from the forested water-
shed during periods of heavy water flow. Smaller particles tend to be
exported during light and heavy rains and subsequent low and high river
flows. Individual storms thus play a large role in controlling particulate
matter export from forested watersheds.
Particulate matter exported from watersheds consists of both organic
and inorganic particles. In the undisturbed, forested watershed, organic
matter constituted a mean of about 39% of all particulate matter exported
during the study. In some years, the proportion was close to 50%, whereas
in other years there was a much higher proportion of inorganic particu-
late matter. Particulate matter from the clearcut watershed without trees
to produce organic particulate matter became increasingly inorganic,
especially toward the end of the study. Higher concentrations at higher
flows late in the study were primarily inorganic, and the researchers con-
cluded that this was caused by increased erosion of inorganic materials
after clearcutting occurred. This process led to changes in losses of par-
ticular elements in comparison to the undisturbed watershed.
Clearcutting had a remarkable effect on the total net loss of all ele-
ments. Even phosphorus (P), for which the absolute values were small,
had a loss rate that was more than ten times higher in the clearcut than
the forested watershed. Phosphorus in inorganic particulate matter in the
undisturbed watershed was 0.009 kg hectare21 and was 0.13 kg
hectare21 in the clearcut watershed. Although neither value is large in
comparison to losses of other elements, the losses are clearly higher in the
clearcut watershed.
Deforestation had a noticeable effect on export pathways of several
elements. These patterns of increase are related to the biogeochemistry
of individual elements, including their occurrence in the rock underlying
 Nutrient Cycling Is a Mechanism of Homeostasis 13

the soil, input from precipitation, ease with which an element cycles and
accumulates in biological systems, and solubility. Biogeochemistry is the
study of the cycles of chemical elements between the living and nonliving
parts of an ecological system. Losses of calcium (Ca) and nitrogen (N)
were extremely large in the clearcut watershed and were mostly lost as
dissolved ions in the water. Magnesium, potassium, and sodium were also
mostly lost from the clearcut watershed as dissolved ions, but losses were
less than calcium and nitrogen losses. Iron was mostly lost in inorganic
particulate matter, and losses of magnesium, potassium, and sodium were
all higher in inorganic particulate matter from the clearcut watershed
than from the undisturbed watershed.
The elements lost as dissolved substances tend to have ionic forms that
dissolve readily in water and can be leached from particulate matter dur-
ing rain events or when particulate matter enters streams. Some elements
that may be found in inorganic material (such as, iron, calcium, magne-
sium, potassium, and sodium) will be found in higher concentrations in
inorganic particles matter, but the latter four can also be leached from
inorganic material and end up as dissolved ions in the water. Nitrogen
may be more likely leached from organic particulate matter.
Nutrients cycle in ecological systems and individual animals and plants
are involved in the cycling of those nutrients. Individual fish maintain
homeostasis of phosphorus, as well as other nutrients and energy, in their
bodies, and this illustrates the organization and energy-dependence of
life. The sum total of all the activities of all the fish in a population exerts
an effect on homeostasis of the ecological system; nutrient cycling is an
emergent property. Abiotic processes also play critical roles in cycling of
nutrients. Movement of nutrients involves feedback mechanisms, which
is a theme in homeostasis. Each nutrient cycle illustrates homeostasis of
ecological systems. At the large and complex ecological system level, pro-
cesses that maintain homeostasis operate over a long time and large spatial
scales, often taking years for a forest to return to homeostasis after a dis-
ruption (such as, clearcutting) that affects ecological system homeostasis.
Nutrients cycle, but other matter also moves through ecological systems.
In the next chapter, what happens to wastes in ecological systems and how
these wastes affect homeostasis will be explored.
14 ECOLOGICAL HOMEOSTASIS

Ethical, Legal, Social Implications: The Gulf of Mexico

Dead Zone Is Related to Increased Nutrient Input
Humans and their activities, including deforestation, agriculture, and
other land use changes, can have a large impact on net export of nutrients
from ecological systems, as the research of Bormann, Likens, and their
colleagues demonstrates. One massive, cumulative alteration of ecologi-
cal systems related to homeostasis of ecological systems is the so-called
Gulf of Mexico dead zone, an area of ocean where dissolved oxygen levels
are very low. This area is near the mouth of the Mississippi River, which
drains about 40% of the continental United States (US), 58% of which
is farmland (Figure 5).
The hypothesized cause of this dead zone is massive nutrient exports
from farmland to the Mississippi River, which then carries the nutrients
all the way to the Gulf of Mexico. Each spring when farmers fertilizer their
fields, excess nutrients are washed into rivers and carried down the Mis-
sissippi River to the Gulf of Mexico. Manure from livestock also contains
nutrients that leach into the river. The nutrients responsible for the dead
zone are nitrogen and phosphorus. Nutrient overloading is part of eutro-
phication, and algal blooms are the result of the nutrient overload. These
excessive nutrient inputs into the ocean can cause toxic algal blooms.
The algal blooms in the Gulf of Mexico tend to occur each sum-
mer along the coast, west of the mouth of the Mississippi River. They
occur in the summer in response to the annual pulse of fertilizer runoff
and west of the mouth because of water currents in the gulf. The result-
ing midsummer dead zone is about 20,000 km2, the second largest such
oxygen-depleted area in the world. Although information gaps still exist,
the overwhelming scientific evidence indicates that algal blooms, driven
by nitrogen loading from the Mississippi River drainage basin, are the
primary source of the organic carbon that decomposes. When produc-
tion increases in an ecological system, organic matter (such as, algal cells)
increases. When this matter sinks to the bottom, bacteria consume it,
and the decay depletes oxygen. Lack of mixing of surface and deep waters
prevents oxygen replenishment. When dissolved oxygen levels drop below
2 mg/L, massive fish kills may result.
The American Midwest is a significant source of food for Americans
as well as people in other countries, and the Gulf of Mexico is a major
 Nutrient Cycling Is a Mechanism of Homeostasis 15

New
Oleans

Sabina Lake Lake Calcasieu

Atchafalaya River
Mississippi River

Terrebonne bay

DO (mg/L)
7

Figure 5 Map of continental US showing the extent of the Mississippi

River basin. The Gulf of Mexico dead zone appears along the coast
west of New Orleans, LA, which is enlarged in the bottom image.
The enlarged area shows dissolved oxygen concentrations for a period
in the summer of 2009. Dots represent sampling sites and the shading
indicates dissolved oxygen at the bottom of the gulf. Areas encircled in
black are below 2 mg O2/L (2 ppm O2).
Source: From http://water.epa.gov/type/watersheds/ named/msbasin/marb_pop1.cfm (US map)
and http://www.noaanews.noaa.gov/ stories2009/20090727_deadzone.html (enlarged area).

source area for the seafood industry. The Midwest produces most of the
over 330 million tons of corn grown by US farmers every year. The US
is the largest exporter of corn with 60% or more of all corn exports com-
ing from the US. Much of the corn is fed to livestock, but corn is also
used to make a variety of food and non-food products, as well as ethanol
for automobile fuel. The Gulf of Mexico supplies 72% of US harvested
shrimp, 66% of harvested oysters, and 16% of commercial fish. Both the
16 ECOLOGICAL HOMEOSTASIS

agricultural and fishing industries have vested economic interests at play,

and their interests are not always compatiblefarmers in Iowa that want
to produce more corn may increase fertilizer applications, which can then
increase nutrient inputs into the Gulf of Mexico, harming the shrimp
industry in Louisiana, for instance.
Homeostasis of both ecological systems is disrupted, first by oversup-
plying nutrients to farms, which results in runoff of nutrients during rains
or irrigation of fields. This is both an ecological and economic loss for the
agricultural systems. A farmer that applies more fertilizer than is needed is
practically throwing his money away. Not only that, he may be indirectly
taking money away from fishermen in the gulf, because if the low oxygen
zone continues or worsens, fishermen and coastal state economies will be
greatly impacted.
There have been many suggestions to remedy the problem. To achieve
the goal of reducing the size and intensity of the Gulf of Mexico dead
zone, scientists recommend a 30% reduction in nitrogen inputs from
Midwest farms. All nitrogen sources should be considered in the strategy,
but because 74% of the nitrate is from agricultural sources and because
56% of the total nitrate originates north of the mouth of the Ohio River,
nitrogen reductions in the upper Midwest will be crucial to effective
implementation of the plan. Farmers will play a key role in success of any
attempt to reduce the size or intensity of the low oxygen dead zone. Farm-
ers can reduce their use of fertilizers or spread their fertilizer applications
over a longer period of time. Ranchers must prevent animal manure from
entering the water. Companies that manufacture products that might
have nutrient waste must monitor and limit the discharge of nutrients
into water. Restoration of wetlands near the mouth of the river would act
as a natural filter, allowing the wetlands to take up the nutrients before
they entered the Gulf. These are a few simple solutions that would drasti-
cally reduce nutrient inputs into the Mississippi River.

Bibliography
Bormann FH, Likens GE, Siccama TG, et al.: The export of nutrients
and recovery of stable conditions following deforestation at Hubbard
Brook, Ecol Monographs 44(3):255277, 1974.
 Nutrient Cycling Is a Mechanism of Homeostasis 17

Bruckner M: The Gulf of Mexico dead zone, Microbial Life Educational

Resources (website): http://serc.carleton.edu/microbelife/topics/dead
zone/. Accessed July 24, 2014.
Kromm C: The Gulf of Mexicos dead zone is among the worlds
largestand corn is one of the culprits, Indy Week (website): http://
www.indyweek.com/indyweek/the-gulf-of-mexicos-dead-zone-is-
among-the-worlds-largestandmdashand-corn-is-one-of-the-culprits/
Content?oid=1520017. Accessed July 24, 2014.
Lewis WM, Jr.: Nitrogen and phosphorus runoff losses from a nutrient-
poor tropical moist forest, Ecology 67(5):12751282, 1986.
Rabalais NN, Turner RE, Scavia D: Beyond science into policy: Gulf of
Mexico hypoxia and the Mississippi River, BioScience 52(2):129142,
2002.
Vanni MJ, Layne CD, Arnott SE: Top-down trophic interactions in
lakes: effects of fish on nutrient dynamics, Ecology 78(1):120, 1997.
 Index
Absorbed photosynthetically active
radiation (APAR), 3031
Accumulators. See Hyperaccumulators
Acid precipitation, 24
Arsenic contamination
in fronds, 2223
in roots, 2223
in soils, 22
Arsenic toxicity, 23
Assimilation, 1, 5, 8
Atmospheric carbon dioxide, 39
Biodiversity, 39, 42
Biogeochemistry, 1213
Bormann, Herbert, 810, 14
Carbon dioxide
concentrations, 27
emissions, 24
increasing atmospheric, 2742
Carbon sink, 27
Carbon source, 27
Climate
defined, 40
change, 27, 32, 34, 35, 39, 40
Cloud water, 3639
Compartments in
ecological systems, 1, 8
Contaminants, 19
Decomposition, 1, 6
Ecological mismatch, 35
Ecological systems
homeostasis of, 27
filter wastes, 1926
Epiphyte mats, 36, 3839
Epiphytes
mortality, 38
viability or growth of, 36
Eutrophication, 14
Free-Air Carbon dioxide Enrichment
(FACE), 2829, 31
Global carbon cycle, 27
Global climate change, 27, 32, 34,
35, 3942
Global temperatures, 27, 32
Global warming, 42
Greenhouse gas emissions, 27
Gross primary production (GPP), 29
Gulf of Mexico dead zone, 14, 15
Homeostasis
in ecological systems, 7, 8, 13, 14,
16, 19, 27
in individual organisms, 19
of nutrients, 8
Hydrologic cycle, 11
Hyperaccumulators, 21, 22
Inorganic materials, 11
Leaf area index (LAI), 31
Leaf mortality, 36, 37
Leaf production, 36, 37
Likens, Gene, 89, 11, 14
Metal percentage in soils, 20
Metal pollution, 19
Monteverde Cloud Forest Reserve
(MCFR), 36
Mutually agreed upon coercion, 26
Nadkarni, Nalini, 3538
National Climatic Data Center, 33
Net primary production (NPP),
2931
Nickel concentrations
in fruits, 21
in plant tissues, 21, 23
in S. acuminate latex, 21
in soil, 21
48 INDEX

Nitrogen fixation, 1, 8 Seston, 3

Norby, Richard, 27, 2932
Nutrient cycling
for ecological systems, 116
deforestation on, 8
Nutrient dynamics, 12
Nutrient exports, 8, 14
Nutrient regeneration, 78
Ocean dumping, 24
Organic debris, 11
Overfishing, 24
Overgrazing on federal lands, 24
Phosphorus budgets, 5, 7
Phosphorus concentration in fish, 34
Phosphorus cycle, 2, 68
Phosphorus movement, 4
Photosynthesis, 1, 27
Pollutants, 19
Respiration, 29
Sebertia acuminate, 20, 21
Seed bank, 38
in epiphyte mats, 38
Solano, Rodrigo, 3538
Solar energy, 27
Summer temperature, 41
Tragedy of the commons, 24
US Environmental Protection
Agency, 26
Vanni, Michael, 27
Water column, phosphorus
concentration in, 2
Water discharge, 9
Watershed treatment, 813
Weather
defined, 40
and climate difference, 4042
Xylem, 23
Zooplankton biomass, 3
 OTHER TITLES IN OUR BIOLOGY
COLLECTION

Behavior and Information Exchangeby Christopher J. Paradise and A. Malcolm Campbell

Cells in Tissuesby Christopher J. Paradise and A. Malcolm Campbell
Ecological Dynamicsby Christopher J. Paradise and A. Malcolm Campbell
Evolution of Interactions in Communitiesby Christopher J. Paradise and
A.Malcolm Campbell
Evolutionary Historyby Christopher J. Paradise and A. Malcolm Campbell
Effects of Genetic and Pathogenic Diseases on Cellsby Christopher J.Paradise and
A.Malcolm Campbell
Information in the Environmentby Christopher J. Paradise and A. Malcolm Campbell
Mechanisms of Evolutionby Christopher J. Paradise and A. Malcolm Campbell
Properties in and of Populationsby Christopher J. Paradise and A. Malcolm Campbell
Variation and Population Geneticsby Christopher J. Paradise and A. Malcolm Campbell
Ecological Interactionsby Christopher J. Paradise and A. Malcolm Campbell
Emergent Properties of Individual Organismsby Christopher J. Paradise and
A. Malcolm Campbell
Organismal Homeostasisby Christopher J. Paradise and A. Malcolm Campbell
Population Homeostasisby Christopher J. Paradise and A. Malcolm Campbell

Announcing Digital Content Crafted by Librarians

Momentum Press offers digital content as authoritative treatments of advanced engineering
topics by leaders in their field. Hosted on ebrary, MP provides practitioners, researchers, faculty,
and students in engineering, science, and industry with innovative electronic content in sensors
and controls engineering, advanced energy engineering, manufacturing, and materials science.

Momentum Press offers library-friendly terms:

perpetual access for a one-time fee
no subscriptions or access fees required
unlimited concurrent usage permitted
downloadable PDFs provided
free MARC records included
free trials

The Momentum Press digital library is very affordable, with no obligation to buy in future
years.
For more information, please visit www.momentumpress.net/library or to set up a trial in the
US, please contact mpsales@globalepress.com.