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Individual organisms contribute to nutrient cycling in ecologi-
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Christopher J. Paradiseis professor of biology and environ-
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A. Malcolm Campbell
Ecological Homeostasis
Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
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
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ISBN-13: 978-1-60650-953-1 (print)

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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).
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
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
= 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
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
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
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
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.
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
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
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.
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
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
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.
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.
CHAPTER 2
Ecological Systems Can

Filter Wastes Like Individual
Organisms
In Chapter 1, it was shown how nutrients are exported from terrestrial

ecological systems as water flows over and through them. Nutrients
exported this way are available to organisms in other systems, and those
organisms may assimilate the nutrients into their cells, tissues, and organs.
The systems may be aquatic ecological systems where the nutrients first
end up, but nutrients may also be exported to the ocean and to other
terrestrial ecological systems during flood events. This movement among
ecological systems is part of nutrient cycles at a larger scale than that of a
single ecological system.
Assimilation of chemicals that are nutrients leads to maintenance of
homeostasis in individual organisms and ecological systems. The property
of ecological system homeostasis emerges from the activities of individual
organisms. There are other chemicals in the environment that may be
harmful. Pollutants, for instance, can disturb homeostasis of organisms
and populations. As a result of certain human activities, contaminants
may enter ecological systems, and those that cause harmful effects on
organisms are termed pollutants. For instance, the processes of mining
and smelting (extracting metals from their ores) expose ecological systems
to contamination by metals. Metals are released by natural processes, such
as erosion, but are released at higher rates in higher concentrations when
mining exposes deep underground rocks to air and water. During smelt-
ing, metals may be released into the air, water, and soil. Soils near smelters
may be highly contaminated; soils near a Middle Ages metallurgical work-
shop were recently found to contain high levels of heavy metal pollution.
Table 1 Percentages of metals in soils from

New Caledonia rich in nickel and from latex.
Approximate ranges of normal percentages are
also shown.
percentage percentage of dry
metal
in soil mass in plant latex
iron 45 0.06
chromium 3 0.004
magnesium 2 0.052
nickel 0.85 25.74
cobalt 0.1 0.007
calcium 0.06 0.52
potassium 0.02 0.15
Source: From Jaffr et al., 1976, in text.
Although metals may remain in ecological systems for hundreds of years,

they may also be transported out or they may be removed or converted by
biological processes as part of ecological system homeostasis.
Organisms growing on or in contaminated soils might adapt to the
high concentrations to which they are exposed. Once it is known that
soil in an area contains high concentrations of a metal, scientists may
investigate the plants in that area for evidence of adaptation. T. Jaffr and
his colleagues studied Sebertia acuminata, a small tree endemic to New
Caledonia, a series of islands in the southwest Pacific (Jaffr et al., 1976).
Endemic species are found only in one particular locality and nowhere
else. There are soils on these islands relatively rich in nickel (Table 1),
which are 10 to 200 times normal concentrations. Nickel in high con-
centrations is toxic to most plants, although is an essential nutrient in
small doses for the processing of nitrogen wastes. Jaffr and his colleagues
examined the content of metals in the latex, a complex liquid consisting
of proteins, starches, sugars, oils, and resins, of S. acuminata trees.
Jaffr and his colleagues examined the content of nickel in various
tissues of these S. acuminata trees. The percentage of dry mass in the latex
was 25.74, in leaves it was 1.17, in bark it was 2.45, and in fruits 0.30. The
scientists also compared the nickel concentrations that they found with
research on several other species, most of which were also known from
New Caledonia. Hybanthus floribundus, from western Australia contained
0.71% nickel in the dry mass of leaves, 0.17% in bark, 0.13% in fruits,
Ecological Systems Can Filter Wastes 21
and 0.48% in flowers. Psychotria douarrei, also from New Caledonia,

contained 3.40% nickel in the dry mass of leaves, 5.24% in bark, 2.30%
in fruits, and 2.40% in flowers. Other species thought to live in similar
environments to S. acuminata contained between 0.6 and 1.45% nickel,
as a percentage of dry mass, in their leaves.
Plants growing in contaminated soils that can tolerate, detoxify, or
store pollutants have an evolutionary advantage on those soils over spe-
cies that do not have those adaptations. Many plant species have been
discovered that have the ability to accumulate high concentrations of
various metals, and are called accumulators or hyperaccumulators. A
hyperaccumulator is a plant that accumulates trace elements from its
environment in concentrations higher than those found in the environ-
ment. Most nickel accumulators and hyperaccumulators belong to spe-
cies found in two genera in New Caledonia. This could be for several
reasons. First, soils rich in nickel could be a selective factor that led to the
adaptation of accumulation and tolerance of nickel. A second reason may
be that because these species are all related, the adaptation could have
been inherited from an ancestral species that evolved to be able to handle
high concentrations of nickel. Third, scientists found these accumulators
here because that is where they looked for them. However, the discovery
of a species in a different genus points to the possibility that accumulation
of toxic metals is widespread among plants living in contaminated soils.
Nickel is found in very high concentrations in S. acuminata latex,
about five times higher than for any other part of any other species.
Calcium and potassium accumulate in S. acuminata, and this makes sense
because they are critical plant nutrients, but it is not clear why nickel would
accumulate. The extremely high nickel concentration in latex may have
evolved as a defense against herbivores, because nickel is toxic to animals
as well as plants. Examining the distribution of nickel in different plant
tissues, one can observe that within any species where multiple tissues were
tested, fruits had the lowest concentration. Fruits provide nutrients and
energy to seeds and are often consumed by animals, which facilitates seed
dispersal. Storing high concentrations of nickel in fruits would not make
much sense for animal dispersal of seeds. High concentrations in leaves and
latex might protect against herbivores, and storage in bark, which is not liv-
ing tissue, would prevent the nickel from coming into contact with living
cells. Hyperaccumulation by plants might be an emergent property of eco-

logical system homeostasis as uptake by some organisms reduces exposure
to other organisms and subsequent disruption of the ecological system.
Arsenic is a heavy metal that is both toxic and carcinogenic. Arsenic
contamination in soils is widespread around the world as a result of min-
ing, withdrawal of groundwater from arsenic-containing rock, burning
of coal, volcanic eruptions, and use of certain pesticides. Lena Ma and
her colleagues studied plants growing on a site in Florida contaminated
with chromated copper arsenate, the compound used in treated lumber,
which is used to control wood pests (Ma et al., 2001). The scientists used
atomic absorption spectroscopy to determine arsenic concentration in
soils and plants. Only one of 14 plants, brake fern (Pteris vittata), showed
large amounts of arsenic (from 3,280 to 4,980 parts per million [ppm]).
The soil samples collected contained between 18.8 and 1,603 ppm arsenic.
Ma and her colleagues collected brake ferns from an uncontaminated site
and grew them in pots of soil spiked with 100 ppm arsenic. The scientists
sampled the roots and fronds, the leaves of a fern, of replicate plants every 2
to 4 weeks. They found very little arsenic in the roots over time, but a dra-
matic increase in arsenic in the fronds, from 0 to over 6,000 ppm in 8 weeks.
The scientists next collected more ferns from an uncontaminated site
and grew them in pots. The pots contained soils from the contaminated
site (400 ppm); uncontaminated soil, which contained a small amount
of arsenic (6 ppm); and potting soil that was spiked with one of three
levels of arsenic (50, 500, and 1,500 ppm as). Fronds were examined
for arsenic after 2 and 6 weeks. In the uncontaminated control soil, Ma
and colleagues found 755 ppm arsenic after 2 weeks and 438 ppm after
6weeks, demonstrating that arsenic rose and then declined when exposed
to very low levels of the toxin. In the arsenic contaminated soil, arsenic in
fronds was 3,525 ppm after 2 weeks and 6,805 ppm after 6 weeks. In the
low arsenic spiked soil they found that fronds contained 5,131 ppm after
2 weeks and 3,215 ppm after 6 weeks. In the medium arsenic spiked soil
they found that fronds contained 7,849 ppm after 2 weeks and 21,290 pm
after 6 weeks. Finally, in the high arsenic spiked soil they found that fronds
contained 15,861 ppm after 2 weeks and 22,630 ppm after 6 weeks.
Brake fern is a strong hyperaccumulator of arsenic. Arsenic concentra-
tions in roots were very low, never exceeding 303 ppm, but concentrations
in fronds quickly reached 6,000 ppm and ultimately climbed to over

7,000 over 20 weeks of growth. Over 90% of the arsenic was concen-
trated in the fronds, yet the roots are the tissues initially exposed to arse-
nic in the soil. One might have expected to observe higher concentrations
in the roots, yet it appears that arsenic is transported to the fronds in
xylem. It is not clear why, although it might have been speculated that the
reasons are similar to the hypothesized reasons behind the distribution of
nickel in plant tissuesdefense against herbivores or storage in cellular
compartments away from cell components or tissues sensitive to arsenic.
Even when exposed to very low concentrations of arsenic, brake
fern accumulated arsenic quickly. Brake ferns growing in soil containing
only about 6 ppm arsenic accumulated over 700 ppm in their fronds in
2 weeks. Non-accumulating plants growing in uncontaminated soil con-
taining trace amounts of arsenic typically contain less than 5 ppm of arse-
nic. Uptake by most plants is low, because roots restrict arsenic uptake,
there is limited translocation from roots to shoots, and plant tissues often
suffer arsenic toxicity at low concentrations. The plants that do not accu-
mulate arsenic would likely not even survive in soils containing 1,500 ppm
arsenic, yet brake fern was tolerant and grew very well in that level of
exposure. Those plants in medium and high arsenic-spiked soils contin-
ued to accumulate arsenic after 6 weeks. It seemed as though the plants
in the high spiked soil were nearing their limit, though, because their rate
of accumulation slowed down and the medium spiked soil plants were
catching up to levels in the high spiked treated plants. At the same time,
plants exposed to low arsenic concentrations, either in the low spiked
treatment or the control soil, lost arsenic after the initial accumulation
in the first 2 weeks. A large input of arsenic may be required to maintain
high levels or to continue increasing; perhaps concentrations that ini-
tially became very high decreased as arsenic was lost back to the soil. Not
enough is currently known about this phenomenon.
Despite this knowledge gap, Ma and her colleagues suggested that
brake fern possesses many attributes that make it a good candidate for use
in reducing arsenic in contaminated soils. In the time since this discovery,
other fern species have been found to hyperaccumulate arsenic. In regards
to use of brake fern to remediate contaminated soil, Ma and her col-
leagues determined that brake fern removes arsenic quickly, even at high
concentrations. In this and other arsenic accumulators, arsenate (AsO43)

has been found to be taken up via phosphate transport systems of the
roots. Once in the roots, it appears that a different mechanism transports
arsenic to the frondsone that does not involve phosphate transport.
Ma and her colleagues determined that brake fern grows well in con-
taminated soils, and it removes arsenic from the soil. In addition, brake
fern accumulates biomass quickly and is easy to grow. Brake fern, and
other plants like it, could be planted in ecological systems where soil is
contaminated with arsenic or other heavy metals. After a brief period of
time, fronds or other aboveground biomass could be harvested, removing
their accumulated contaminants from the ecological system. This would
help restore homeostasis in the affected ecological system. This is just one
example of how individual organisms in an ecological system play a role
in maintaining ecological system homeostasis.
Ethical, Legal, Social Implications:

Pollution Is a Tragedy of the Commons
Ecological systems have the ability to deal with pollutants released into the
environment, but pollutants may overwhelm a system and disrupt homeo-
stasis. The extraction of resources, including metals from mining opera-
tions, and the dumping of waste products resulting from the mining are
two examples of what has been called the tragedy of the commons. The
commons refers to a limited resource shared among a group of people. The
limited resource might be owned by several to many people or not owned
by anyone. Anyone in the group has use of the resource. An example of
the former is a well that is used for drinking water and owned by a village,
and an example of the latter is clean air. Aristotle noted that what is com-
mon to the greatest number has the least care bestowed upon it. A well
used for drinking water might suffer from overexploitation of water, and
clean air might be compromised as pollutants are pumped into it. In both
situations, if there are too many people using the resource and use of the
resource is not managed, then the resource will be destroyed or depleted.
In a broad sense, the tragedy of the commons can be seen in many mod-
ern environmental problems, including overgrazing on federal lands, acid
precipitation, ocean dumping, carbon dioxide emissions, and overfishing.
The tragedy is the dilemma that arises as individuals, acting indepen-

dently and rationally in their own self-interest to maximize their own
short-term gain, deplete the shared resource even when it is clear that it is
not in anyones long-term interest. This behavior can cause long-term harm
to the environment, other individuals, and ultimately oneself. The more
people who use the resource, the faster it is destroyed or depleted; thus
the tragedy of the commons is related to population size. In many human
societies, the environment has been treated as a free dumping ground for
pollutants. Humans and their corporations have attempted to maximize
short-term economic gains by unloading waste products into the environ-
ment at little or no cost. Viewed as an open access resource, free to use by
anyone, the environment was not factored in to the cost of doing business.
The contamination of soils by heavy metals discussed in this chapter
is a tragedy of the commons in the sense that clean, fertile soil and clean
groundwater, both of which suffer, are limited resources. These resources
are common property in that they belong to all of us or to none of us. A
miner might own the land on which metals are dumped, but the contam-
ination does not stay within property boundaries. Similarly, metals that
leach out of treated lumber may contaminate the environment. The
contaminant may be transported in the air or water to other locations,
crossing property, state, and international boundaries.
Once it is recognized that a particular environmental issue is a tragedy
of the commons, scientists and policy-makers might investigate the cause
and possible solutions. Various solutions to the tragedy of the commons
have been proposed, and they have been debated among economists,
policy-makers and environmentalists for many years. Some argue that
tragedy can be avoided by limiting the size of the human population.
Alternatively, some form of authority may be needed to regulate or limit
the amount of an extracted resource or the amount of pollution that can
be emitted by a user. The authority, possibly a government, might use a
permit system, tax incentives, or fines to regulate the resource. Another
solution would be a cooperative agreement among resource users to facili-
tate resource conservation in the interest of mutual benefit.
Still others have argued that there can be no technical solution, that is,
one that requires only a change in the techniques or methods that might
reduce depletion or destruction of a resource. These people contend that
without a change in human values, human livelihoods, or ideas of moral-

ity, these technical solutions will not work. For example, human society
may decide that unlimited pollution is no longer morally acceptable, and
only in this way will technical solutions be accepted and adopted. Fur-
ther, changes in values can lead to changes in resource use, where mutu-
ally agreed upon access restrictions or incentives for users to invest in
resources instead of depleting them might be implemented. However,
others argue that as long as humans retain a human-centered ethical sys-
tem, where humans are the primary concern, then human overpopulation
and overconsumption are constant threats to commonly held resources.
If society agrees that some actions are no longer allowed by mutual
agreement, society may then impose fines or prison terms on those who
violate that agreement. This has been called mutually agreed upon coer-
cion. In the US there are restrictions on what can be put into the air,
and the US Environmental Protection Agency regulates the amount of
air pollutants. Failure to comply with the regulations can lead to fines or
prison sentences. Even though these laws arose in a human-centered ethi-
cal system, there is recognition among many humans that the decline of
ecological systems, although not the concern of a human-centered ethics,
ultimately harms human health and human societies.
Bibliography
De Young R: Tragedy of the commons. In Alexander DE, Fairbridge RW,
editors: Encyclopedia of environmental science, Hingham, MA, 1999,
Kluwer Academic Publishers.
De Young R: Tragedy of the Commons (website): http://www-personal
.umich.edu/~rdeyoung/tragedy.html. Accessed July 24, 2014.
Hardin G: The tragedy of the commons, Science 162:12431248, 1968.
Jaffr T, Brooks RR, Lee J, et al.: Sebertia acuminata: a hyperaccumulator
of nickel from New Caledonia, Science 193(4253):579580, 1976.
Ma LQ, Komar KM, Tu C, et al.: A fern that hyperaccumulates arsenic,
Nature 409:579, 2001.
Wang J, Zhao F-J, Meharg AA, et al.: Mechanisms of arsenic hyperac-
cumulation in Pteris vittata. Uptake kinetics, interactions with phos-
phate, and arsenic speciation, Plant Physiology 130:15521561, 2002.
CHAPTER 3
Increasing Atmospheric
Carbon Dioxide Can Disrupt
Ecological Systems
Mean global temperatures are rising, due in part to human activities that
have led to increases in levels of CO2 and other greenhouse gases. Yet
global efforts to reduce greenhouse gas emissions have stalled in part
because of how climate change science is often manipulated by policy-
makers. Carbon dioxide concentrations can be altered by photosynthesis.
In examining the efficiency of energy transfer in ecological systems, it can
be quickly seen how energy flow and the carbon cycle are linked. Ecologi-
cal systems affect and are affected by the global carbon cycle, greenhouse
gases, and solar energy. These ideas will be explored further in this chapter,
with a focus on how increasing greenhouse gases, and the climate changes
that result, affect homeostasis of ecological systems.
Trees photosynthesizing in Brazils tropical rainforests affect glaciers of
Greenland through their effects on atmospheric CO2 concentrations. A
rainforest actively photosynthesizing acts as a carbon sink, whereas one
that is cut down by humans could be a carbon source. A carbon sink
is a reservoir that accumulates and stores carbon-containing chemicals,
while a carbon source is a reservoir that releases carbon-containing chemi-
cals. Other carbon sources are also related to human activities. Besides
deforestation, forest ecological systems may be affected by changing envi-
ronmental factors, including rising CO2 and temperatures, and changes
in precipitation patterns associated with global climate change. Richard
Norby and colleagues from several research institutions compiled data
from experiments on the effects of elevated CO2 on growth and produc-
tivity of forests (Norby et al., 2005).
FACE plots
loblolly
pine forest
Figure 6 The FACE experimental site at the Duke University Forest.

FACE plots are located in a primarily loblolly pine forest, including
many deciduous trees. The photo is taken in autumn, and deciduous
trees can be seen amongst the pines. Each FACE plot consists of an
array of vertical pipes surrounding the plot and CO2 sensors inside of
the plot.
Source: From https://www.bnl.gov/ bnlweb/pubaf/pr/1999/bnlpr051399.html.
The scientists involved in this study work at several sites in forests in

the US and Europe where Free-Air Carbon dioxide Enrichment (FACE)
devices have been set up. FACE devices are made up of tall vertical pipes
placed in a circle around an experimental plot, which are up to 30 meters
in diameter (Figure 6). The pipes emit CO2-enriched air toward the inte-
rior of the plot to maintain a higher concentration of CO2. Sensors in the
plots provide feedback to computers, which monitor and adjust the out-
put from the pipes. Some arrays serve as control plots and pump ambi-
ent air into the plot, whereas other control plots have no arrays. Various
responses of the plants, including growth rates and primary productivity,
inside the plots are measured over long periods of time.
FACE experiments are expensive, yet they are critical because measur-
ing the effects of elevated CO2 in ecological systems can better estimate
how plant growth and forest communities will change as CO2 concentra-
tions rise. FACE experiments also allow the effect of elevated CO2 on
INCREASING CARBON DIOXIDE CAN DISRUPT ECOLOGICAL SYSTEMS 29
tall, long-lived plants (such as, trees) to be measured. Norby and his col-
leagues analyzed net primary production (NPP) data from four long-
term FACE experiments, in Wisconsin, North Carolina, Tennessee, and
Italy. The goal was to assess how increased levels of CO2 over long periods
of time affect forest ecological systems and through that determine how
the global carbon cycle might be affected, as forests dominate global NPP.
FACE experiments are all conducted in temperate zone forests, yet
the forests represent a broad range of productivity, climatic conditions,
and diversity of tree species. Mean annual precipitation ranged from
810 (Wisconsin) to 1390 (Tennessee) mm/yr and mean annual tempera-
ture ranged from 4.9 (Wisconsin) to 15.5 (North Carolina). Growing
seasons ranged from 150 (Wisconsin) to 247 (Italy). The North Carolina
FACE was dominated by loblolly pine, Wisconsin by aspen, maple, and
birch, Tennessee by sweetgum, and Italy by poplar.
For the experiments, the average atmospheric carbon dioxide concen-
tration ([CO2]) in the ambient CO2 plots across all of the years assessed
was 376 ppm, whereas the elevated CO2 FACE plots had an average
[CO2], across all experiments and sites, of 550 ppm, which is a level cho-
sen based on projections of atmospheric [CO2] anticipated for 2050.
The FACE approach permits the study of processes under elevated
CO2 that was previously impossible or intractable in forest ecological sys-
tems. As with any experiment, experiments in ecological systems require
adequate controls, and the FACE experiments included two controls. The
first control was the use of FACE arrays that blew ambient, or current,
CO2 into plots. This control is necessary to test for the effects of elevated
carbon dioxide relative to current levels. The second control of plots mon-
itored over time, but with no arrays, is necessary to insure that plots with
FACE arrays do not suffer any effects due to the array itself, regardless of
the air that is pumped into the plot. To test for any array effect, scientists
would compare the two kinds of control plots, with and without arrays.
Estimates of annual increases in biomass of plant tissues are an esti-
mate of NPP, equal to gross primary production (GPP) minus respira-
tion. The scientists made estimates of biomass increases of stem wood
and large woody roots using periodic measurements of the diameter of
trees in different plots. Leaf tissues were harvested to estimate biomass
of leaves. The scientists estimated growth increases of small root biomass
from known growth rates for particular species and through experiments
where growth of small roots was monitored over time. The scientists used
these estimates of individual stems, leaves, and roots to estimate biomass
increases for entire plots. The scientists collected and estimated the pro-
duction of leaf litter using litter baskets, which collected leaves over a
certain period of time.
NPP was expressed as grams of carbon fixed per square meter of land
per year (g C m22 yr21). It was calculated as annual carbon increases
in wood, leaf, and root tissue, plus the major inputs to detritus, leaf litter,
and fine root turnover. The biomass estimates were converted to carbon
units by using the known carbon content of different tissues, which were
found to vary between 0.4 and 0.5. Thus the scientists converted accumu-
lated biomass to carbon dioxide fixed into organic carbon, which became
their estimate of NPP.
Norby and his colleagues analyzed NPP based on all years and plots
for which they had data and for which the tree canopy was fully devel-
oped; in other words, they excluded years and plots with younger, rapidly
growing trees. They compared NPP in plots with elevated CO2 to that in
plots with current CO2 for each tree species or tree species combination.
The scientists wanted to understand the response of forests with fully
developed canopies, which they reasoned would be informative of global
forest responses.
Norby and his colleagues also measured absorbed photosyntheti-
cally active radiation (APAR, or the light energy absorbed by the can-
opy) by taking the difference of the amount of light energy at the top
of the canopy and that striking the forest floor. These values were then
integrated over the entire growing season to yield the total light energy
absorbed per unit area per year for each tree species combination. The sci-
entists compared APAR in plots with elevated CO2 to that in plots with
current CO2. If elevated CO2 plots were absorbing more light than in the
current CO2 plots, then a best-fit line for all plots would be greater than
a 1:1 relationship line.
Norby and his colleagues found a strong linear relationship between
NPP in elevated CO2 and that in current CO2 plots that rose at a rate
significantly greater than a slope of 1 (P <0.0001). The conclusion of the
scientists was that there was a significant effect of elevated CO2 on NPP,
and an estimated 23% increase in forest NPP would result from atmo-
spheric CO2 increases to 550 ppm over the next few decades. However,
APAR in elevated CO2 plots was not different from APAR in current
CO2 plots. When all FACE plots were graphed, the slope of the best-fit
line was not significantly different from one. Thus, APAR did not increase
faster in elevated CO2 plots relative to current CO2 plots, as NPP did.
Norby and his colleagues compared the NPP response to any potential
change in the light energy absorbed by the plants to determine whether
changes in NPP were related to changes in APAR. They examined this by
standardizing the NPP and APAR values to the maximum values observed
for each plot. The scientists then calculated the ratio of the change in
APAR between elevated and current CO2 plots to the change in NPP
between elevated and current CO2 (APAR/NPP). They plotted this
against the maximum leaf area index (LAI) for each year. LAI is the ratio
of total upper leaf surface area divided by land surface area, and typically
ranges from 0 for bare ground to 7 for a dense forest.
The scientists might have predicted that APAR would be higher in
elevated CO2, because higher NPP should lead to more leaves to absorb
more light energy and might show the same trend across FACE plots as
NPP did. Because this did not occur, Norby and his colleagues hypoth-
esized that higher light-use efficiency caused the increased productivity.
They estimated this by calculating the fraction of the standardized gain
in NPP that was attributable to a gain in APAR. If the standardized ratio
of the change in APAR to the change in NPP is equal to one, then both
parameters are increasing equally, and the increase in NPP is due to an
increase in APAR. However, if the ratio is close to zero, then increases in
NPP are due to higher light-use efficiency.
Norby and his colleagues found that the change in APAR from cur-
rent to elevated CO2 in dense forests is close to zero. In forests with dense
canopies, very little of the gain in NPP was associated with increased
absorbed light energy, whereas in the forest stands with lower peak LAIs,
60% to 90% of the gain in NPP with CO2 fertilization was associated
with increased absorbed light energy. When the canopy is sparse and CO2
is elevated, plants can respond by absorbing more light energy, whereas
when the canopy is dense, the plants in elevated CO2 are converting more
light energy to organic matter per unit energy captured.
Norby and his colleagues presented evidence that suggests a feedback

between the biosphere and atmosphere that involves the carbon cycle.
This is an example of ecological system homeostasis, although the change
in the environmental conditions (increased CO2) may actually cause the
ecological system to move to a new state. A sustained or large change in
an environmental condition could push an ecological system to a new
set point, which is the condition at which a biological system attempts
to maintain itself through feedback mechanisms. Coincident to the sus-
tained change in CO2 concentrations that are occurring globally is the
increased global mean temperature that has been documented. Scientists
predict that by 2100, the Earths mean global temperature could be 1.5o
to 5.8o C warmer than it is now, although the distribution of temperature
changes varies globally. Far northern latitudes have already experienced a
change of between 0.6o to 1.0o C over the last 100 years, which is a change
that is greater in magnitude than for the entire northern hemisphere.
Some changes associated with temperature increases may be detect-
able over the decades-long time span during which temperatures have
been increasing. Nina Bradley and her colleagues investigated pheno-
logical changes in springtime events on a Wisconsin farm over a 61-year
period (Bradley et al., 1999). Phenological changes are periodic biologi-
cal phenomena that occur seasonally or annually. Each year in temperate
latitudes ecological systems progress through a series of events, such as
flowering and fruiting of plants, migration of animals, and reproductive
seasons for a variety of species. This cycling of annual events has been
used by naturalists to mark the passage of the seasons. For instance, in
the northeastern US, the return of the migratory American robin is often
taken as a sign of spring. Scientists have predicted that global climate
change and the associated temperature increase would alter the timing of
some of these events.
Bradley and her colleagues had a rare opportunity to examine long-
term phenological data collected at a farm in southern Wisconsin dur-
ing two intervals by two different scientists. From 1936 to 1947, the
famous biologist and naturalist, Aldo Leopold, maintained records of
spring events on this farm. Twenty-nine years later, his daughter, Nina
Leopold Bradley, began maintaining similar records, which she contin-
ued for the next 22 years (1976 to 1998). The records spanned a total of
61years and included 74 annual biological events, such as arrival dates for
migratory birds and dates of first bloom of spring flowers. The scientists
estimated that recorded events were accurate to within 4 days during the
first 11-year period and 2 days during the later 22-year interval. Events
may be missed due to the recorder not being on the farm constantly, poor
weather conditions, or a missed event, which is later recorded on a dif-
ferent date.
Although many records were kept, the researchers limited their analy-
sis to biological events that typically occur before the end of June. They
used only events for which they had at least six annual records in each of
the two recording intervals. Fifty-five biological events fit those criteria.
The scientists next plotted the day of the year (the ordinal date, or num-
ber of days since January 1st for any year) that an event occurred over all
years for which they had data. They used regression analysis and fit best-fit
lines to the data to determine whether the day of the year changed as the
years progressed. The slope of the line, if statistically significant, would
provide information on the rate of change occurring over time.
The records of biological events on the farm collected by Leopold
and Bradley did not include consistent temperature data. As a proxy for
temperature change over the years, Bradley and her colleagues used the
ordinal date of ice-melt in a nearby lake and studied it using the same
regression analysis used for biological events on the farm. A proxy is a
variable that stands in for another variable. Ice-melt data were obtained
from the Wisconsin State Climatology Office. Most of the melt-dates
occurred in March, so the scientists also plotted the ordinal melt dates
against the mean March temperature, which was data they obtained from
the National Climatic Data Center. Both of these regressions had slopes
that were significantly less than zero (0.123 day per year and 2.72 days
per o C of March mean temperature), indicating that ice melt was occur-
ring earlier and temperatures were becoming warmer as time progressed.
There was some overlap in the ranges of slopes that are significantly
different from zero and those that are not. A slope of 0.231 was found to
be statistically significant for the arrival of cowbirds; they arrived on aver-
age 0.231 days earlier each year over the 61 years of recordkeeping. The
slope of 0.244 is a larger value, yet it was not statistically significant. This
indicates that the latter data probably has more scatter around the best-fit
line, which decreases the probability that the slope is statistically different
from a slope of zero. Large slopes are not always statistically significant,
and small slopes may be significantly different from zero if the scatter
around the line is minimal, as it is for the rose-breasted grosbeak.
If the day of first blooming of a flower species or the day when first
migrants of a bird species return, varies from year to year that indicates
that some environmental factor that also varies from year to year may be
controlling blooming or migration. This will cause scatter in a plot of
ordinal day of year against year. Although temperature has been increas-
ing over the past decades, it does vary from year to year on any particular
ordinal date, and these two observations can cause the patterns observed
by the scientists. If the return is less variable, then something that does
not vary from year to year on a particular day, such as the length of day-
light, might be the cue that causes blooming or migration. If this is the
case, not only would there be less variation from year to year, but the
slope of the line would be zero. There may still be some variation caused
by individuals who vary in their genomes responding differently to the
same invariant signal, but one might predict less variation than if the cue
were changing over time.
The scientists documented 18 out of 55 (32.7%) biological events with
significantly earlier springtime occurrence from about a week to almost a
month earlier over 61 years. Fourteen out of 55 (25.4%) events occur earlier,
although not significantly, according to Bradley and her colleagues. Over
the same period, the surface temperatures of the planet have warmed. These
trends should suggest a connection between first occurrence of biological
events and global climate change, and the ice melt data serve as a useful
substitute for the lack of temperature data from the farm. Ice-melt is
occurring earlier, Wisconsin spring temperatures are rising, and species are
responding to warming by migrating or blooming earlier in the year. This is a
finding that other scientists have found elsewhere for other biological events.
Less than half of the events on the farm did not change when they
first occur, and one occurs significantly later. Many biological events have
been found to be controlled by photoperiod. Among the species that do
not show adaptability to climate change are the species in which seasonal
changes are regulated by photoperiod or possibly some other genetic reg-
ulatory system.
There are at least two implications to the fact that some events are
occurring earlier, some are not changing, and some are occurring later.
First, some species may not be as adaptable as others, and these species
may not survive or may experience greater stress during climate change.
Second, for pairs of species that interact with one another where one
changes its phenological patterns and the other does not, an ecological
mismatch may occur. An ecological mismatch is a change in one species,
but not in an interacting species, during climate change. If a migrating
bird arrives at its breeding ground earlier as the climate warms and its
main food source (critical for rearing offspring) are insects that have not
changed their hatching time, the birds may fledge fewer offspring, lead-
ing to a lower population size over time. Altered interactions could cause
a disruption of ecological system homeostasis as species that interacted
previously do not continue to interact as strongly when their activities no
longer occur in sync.
Global climate change disrupts ecological systems through changes in
temperature, but as temperatures increase, other aspects of climate also
change. For instance, tropical equatorial forests at high altitudes could
be experiencing more dramatic climate change than tropical forests close
to sea level. These tropical cloud forests, or high elevation moist forest
with persistent clouds in the tree canopy, contain species adapted to live
in conditions where clouds persistently form in tree canopies. In other
words, they live in very moist conditions with near constant clouds and
mist. Theoretical models predict that even a slight rise in greenhouse gas
concentrations will increase the altitude at which clouds form in tropical
mountains, up to 200 meters higher. Global climate change has warmed
the sea surface, and this leads to increasing evaporation from oceans. The
water vapor climbs to higher altitudes, as warm air rises, and when the
vapor cools and condenses, latent heat is released. The regional cycling of
water will then be altered in such a way that the amount and frequency of
inputs of mist (low-intensity wind-blown precipitation) will be reduced.
Water in clouds that condenses on vegetation will also be reduced.
Nalini Nadkarni and Rodrigo Solano studied the responses of epi-
phytes in cloud forest to the predicted drier conditions under global cli-
mate change (Nadkarni and Solano, 2002). Cloud forest epiphytes dwell
in the canopy and have no connections to the ground. These plants play
a critical role in cloud forests, because they retain mist and cloud water
and capture both water and nutrients that are cycled throughout the rest
of the forest ecological system.
The researchers set up their study in the Monteverde Cloud For-
est Reserve (MCFR) in Costa Rica, where total annual precipitation is
between 2,000 to 2,400 mm. The canopy there is exposed to frequent
wind and mist throughout much of the year. The researchers used an
existing gradient of cloud water and mist across an elevation gradient,
taking advantage of the fact that moisture conditions at lower elevations
are similar to what is predicted for high elevations under future global
climate change. Because epiphytes often grow together in mats on tree
limbs, the researchers could transplant pieces of epiphyte mats from one
tree limb to another. These mats consist of interwoven roots, bark, and
decaying organic matter up to 25 centimeters thick, and they support
many epiphyte species. The mats can be peeled off and rolled up like a
carpet, so plants and the canopy soil remain intact.
Nadkarni and Solano first tested for any effects of transplantation by
transplanting mats within the upper site and found no statistically signifi-
cant differences in leaf production or leaf or plant mortality between con-
trol and transplanted mats. They concluded that transplantation does not
affect the viability or growth of epiphytes. This allowed them to proceed
to transplant mats to other sites. Differences found in transplant experi-
ments should then be due to site differences, not transplantation effects.
For the transplant experiment, Nadkarni and Solano established
three study sites along an elevation gradient of 150 meters. Cloud water
progressively declined during the dry season from the high elevation site
(1,480 m) to the two transplant sites, the midpoint of the gradient site
(1,410 m) and the low elevation site (1,330 m).
The researchers moved epiphytes from the high elevation site (where
cloud water is frequent throughout the dry season) to trees growing at
the two lower elevation sites, both of which had longer dry seasons. The
researchers climbed trees in the high elevation site, cut 75 centimeter
epiphyte mats, transported them to mid and low elevation sites, and tied
them to branches of three to four trees at those sites. Control mats were
set up at the high elevation site.
Nadkarni and Solano monitored leaves of plants from four representa-

tive epiphyte genera. They counted all of the leaves for each plant belong-
ing to each of the four species (Guzmania pachystylis, a bromeliad; Clusia
sp., a woody shrub; Peperomia sp., a climbing herb, and Elaphoglossom sp.,
a fern). They continued this experiment for 18 months after transplanta-
tion. The transplant experiment was performed twice; once starting in the
dry season (January) and once starting in the wet season (June).
Each month, new leaves were marked with paint, and leaf production
was calculated as the number of new leaves that appeared during each
sampling interval. Leaf mortality was calculated as the number of leaves
missing from the previous sampling. These values were then expressed
as percentage leaf change, based on the initial number of leaves. Finally,
Nadkarni and Solano determined plant longevity as the number of
months the entire plant survived from the beginning of the study.
Nadkarni and Solano found large and consistent effects of elevation.
As one might expect, the epiphytes left in the high elevation site had
positive leaf production and lived for a long time. Epiphytes transplanted
at the beginning of the dry season to mid and low elevation sites did not
produce leaves, and they had a decrease in the percentage of new leaves,
meaning more leaves died than grew. This effect was consistent across all
epiphyte species. The effects on epiphytes transplanted at the beginning
of the wet season were more variable, but only three of four species at the
low elevation site had significantly less leaf production than controls at
the high elevation site. All four species, when transplanted at the begin-
ning of the dry season, had significantly reduced longevity in the mid
and low elevation sites relative to the high elevation site. Further, the
herb and the fern had lower longevity in the low relative the mid eleva-
tion site. When transplanted in the wet season, the scientists found that
only the bromeliad had significantly lower longevity and only at the low
elevation site.
Nadkarni and Solano measured the amount of cloud water input and
moisture content of the epiphyte mat substrates during 4 months of the
dry season at each site. They collected cloud water from collectors that
they checked every 3 to 4 days and determined relative moisture from
small artificial blocks that absorbed moisture from the mats. Sensors were
embedded in the blocks, and these could be checked from the ground
with wires that ran from the ground up to each block. Moisture levels
were checked every 3 to 4 days.
Measurements of canopy moisture input and moisture content of
epiphytes were important for two reasons. It linked elevations to mois-
ture, and it confirmed that epiphyte mats received progressively less water
across the elevation gradient. The elevation gradient was an adequate
substitute for moisture inputs. The measurements revealed that the low
elevation cloud water collectors received less than one-third of the total
water collected at the high elevation site with an intermediate amount in
the mid elevation site. The moisture content of the epiphyte mats paral-
leled the inputs of water. These results allowed the researchers to conclude
that moisture differences played a part in the responses of transplanted
epiphytes relative to control epiphytes.
Epiphyte mats contain a diverse seed bank, which is the natural stor-
age of seeds within most ecosystems, containing both ground and epi-
phytic species. Nadkarni and Solano set up a greenhouse experiment to
determine what would happen in mats if existing epiphytes at high eleva-
tion sites began to suffer higher mortality. They collected 24 epiphyte
mats from trees in the high elevation forest. The researchers pruned off all
of the stems of epiphytes from half of the mats. All mats were placed in a
greenhouse with an automatic misting system so that mats were watered
twice per day. Temperature conditions were kept close to the temperature
of the high elevation site. Over the course of 1 year, the researchers identi-
fied the species of each emerging seedling, measured its height, and tallied
the numbers of each species (Table 2).
Table 2 Results of greenhouse experiment on

seed banks in epiphyte mats. An asterisk after
the variable indicates that pruned mats were
statistically significantly different from the control,
unpruned mats.
variable pruned unpruned
seedling abundance/mat* 127.6 27.1
total number of species* 37 25
percentage of terrestrial species* 90.4 0.9
Source: From Nadkarni and Solano, 2002.

Exposure to conditions with less cloud water, which is a prediction for

tropical cloud forests under global climate change, can lead to adverse effects
on growth and survival of epiphytes. If individual epiphytes die or grow
slowly, populations of those species will also suffer slower growth, shrinking
in size and possibly going extinct. Loss of dominant epiphytes from the cloud
forest canopy might favor a change in community composition in those
ecological systems. Change in the composition of species in a community
is a disruption of ecological system homeostasis. The greenhouse experiment
conducted by the scientist confirmed that loss of epiphytes from mats leads
to emergence of a high density of seedlings, many of which are not epiphytes,
but more typically grow on the forest floor, after removal of epiphytes.
If epiphytes disappear under drier conditions, terrestrial plants will
temporarily dominate the epiphyte mats. However, those plants are not
adapted to living in canopies; and once they die out, epiphyte mats will
become devoid of plants, and the mats will eventually fall apart, because
they are not being replenished by cloud water, nutrients, and organic
matter normally captured and deposited by epiphytes. Only the most
desiccation tolerant epiphytes will survive in those conditions, leading to
a lower diversity and abundance of plants in tropical cloud forests. Epi-
phytes capture water and nutrients that become available to other plants,
and they produce the mats that provide habitat for many animals. Epi-
phyte communities contribute to the biodiversity of tropical ecological
systems, and their loss affects homeostasis.
Beyond epiphytes, organisms in tropical cloud forests, also adapted to
moist conditions, may suffer as global climate changesfurther disrupt-
ing ecological system homeostasis. Disappearing rainforests, as suggested
at the beginning of this section, could be a source of carbon dioxide to the
atmosphere. If so, increasing carbon dioxide in the atmosphere leading to
global climate change and the loss of tropical trees would cause further
increases in atmospheric carbon dioxide, in a positive feedback. Most
feedback mechanisms in homeostasis are negative feedback mechanisms,
correcting or reversing a change in state back to the original state. A posi-
tive feedback mechanism moves the system further away from the original
state, leading to a large disruption in homeostasis. This could lead to loss
of entire rainforests, and it has been suggested that rainforests are a key fac-
tor in maintaining ecological system homeostasis for the entire biosphere.
Ethical, Legal, Social Implications: There Is a

Difference Between Weather and Climate
Weather is defined as a particular event related to the state of the atmo-
sphere at a given time and place with respect to temperature, moisture,
wind, and air pressure, and climate is the long-term average temperature
and precipitation for a given time and place. Weather also includes sun-
shine, clouds, winds, floods, blizzards, thunderstorms, heat waves, and
more. Weather may change from minute-to-minute, hour-to-hour, day-to-
day, and season-to-season. Aspects of climate also include humidity, sun-
shine, wind velocity, and events such as fog and frostall measured over a
long time period. Changes in climate tend to be gradual and are predicted
using averaged weather statistics over long periods of time and space.
Besides the question in the title of this section, one might also ask,
What is the difference between an event and a trend? The answer to both
questions is similar. Weather is the event, whereas climate portrays the
trend. Climate is what is expected based on long-term trends (hot North
Carolina summers) and weather is what is experienced on a day-to-day basis
(a cool day in the middle of July). Considering that the relationship between
weather and climate is similar to considering individual data points around
a trend line. As seen in many datasets that contain a trend line, very few
data points lie on the trend line. Picking out the trend in a set of data is like
picking out a signal from amidst noise. Scattered data are the weather noise,
and the long-term trend is the climate signal. Highly variable weather can
make detection of a signal and forecast of future trends difficult.
There are many factors that contribute to the response of any biological
system to any independent variable. The same is true for weather events,
which is why they are difficult to predict beyond the very near future.
There is extensive evidence that human activities that emit greenhouse
gases into the atmosphere have resulted in both climate and weather alter-
ation from the local to the global levels. Global climate change includes
the slow and steady increase in global mean temperatures. However, it
can also include greater variation in weather events. Climate change
is expected to affect the frequency of extreme weather events, such as
drought, extreme temperatures, flooding, and severe storms. An unusu-
ally cool summer is often used to deny the existence of global climate
deviation from 30 year (19611990) mean temperature (C)

2
3
1880 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010
year
Figure 7 Anomalies for mean summer temperatures (July, August,

and September) for Charlotte, NC. Anomalies are deviations from the
30-year mean (1961 to 1990). The equation for the best-fit trend line
is o C deviation 5 (0.0040 3 year) 2 7.64. The year circled in gray
is referenced in the text.
Source: Data from the National Oceanic and Atmospheric Administration (http://www.erh
.noaa.gov/gsp/climate/cltcli.htm). C. Paradise.
change. However, one weather event cannot be used to deny the existence
or support the reversal of a long-term climate trend (Figure 7). The fig-
ure shows a summer cooler than the average in Charlotte, NC, in 1968,
which is circled in blue. That is an example of a weather event.
Figure 7 also shows the deviation of mean summer temperature for
each year from a 30-year mean summer temperature for Charlotte, NC.
The 30-year mean is from 1961 to 1990, a standard reference range often
used in such studies. The data show a lot of variation, but a best-fit line
indicates that the average deviation in 1880 was less than zero (less than
the 1961 to 1990 average, which is 24.51o C) to well over zero by 2010.
The deviations become more positive over time. The temperature pre-
dicted by the best-fit line for 1880 is 24.35 and that for 2010 is 24.87,
which means that average temperatures have increased by 0.52o C over
the 130-year time period, which is in line with global data (an estimated
increase of 0.76 C (0.19 C)). This is an example of climate.
The effects of global climate change are serious and may pose seri-
ous threats to many aspects of human civilization and ecological systems,
including biodiversity, ecosystem services, availability of natural resources,
food and fiber production, economic development, and human health.
Using individual weather events as evidence for climate change is not
good science. Cold temperatures are not proof that there is no such thing
as global warming.
Bibliography
Bradley NL, Leopold AC, Ross J, et al.: Phenological changes reflect cli-
mate change in Wisconsin, Proc Nat Acad Sci 96:97019704, 1999.
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Masek JG: Stability of boreal forest stands during recent climate change:
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Nadkarni NM, Solano R: Potential effects of climate change on canopy
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Conclusion
Large, complex systems can maintain homeostasis or have it disrupted by
external forces. Even ecological systems can maintain or return to constant
internal conditions in the face of a varying external environment, which is
a definition of homeostasis. Ecological systems, like individual organisms,
have internal conditions and are connected to other biological systems,
meeting the conditions of the definition. Throughout several books in
this series, the concept of homeostasis at various levels of the biological
hierarchy has been explored, and with this book, the examination is com-
plete. As with biological systems at lower levels of the hierarchy, such as
populations, individuals, and cells, ecological systems use negative feed-
back mechanisms to maintain homeostasis. The processes investigated
that regulate ecological systems are time- and energy-dependent, and the
processes in large ecological systems not unique to ecological systems. In
fact, many of the processes are used by biological systems at lower levels,
and those processes affect properties and conditions of ecological systems,
an example of emergent properties.
Glossary
assimilation. Assimilation is the process of absorbing nutrients and incorporating
them into the body.
biogeochemistry. The study of the cycles of chemical elements between the living
and nonliving parts of an ecological system.
biomass. The total amount of organic matter in an organism, population, or
other ecological system after water is removed.
carbon sink. A reservoir that accumulates and stores carbon-containing chemicals.
carbon source. Any natural or human source of carbon dioxide emissions, includ-
ing decomposition, ocean release and respiration (natural) and cement produc-
tion, deforestation, or burning fossil fuels (human).
compartments. A part or space into which an ecological system is subdivided.
contaminants. A potentially harmful or undesirable substance that makes a place
or substance impure when it is added, usually of human origin.
decomposition. The decay into constituent parts by physical, chemical, or bio-
logical processes.
discharge. The volume of water in a river flowing past a point during an interval
of time.
ecological mismatch. A change in one species, but not in an interacting species,
during climate change.
ecological system. An ecological community together with the abiotic environ-
ment, usually considered to function as a unit.
enclosure. A device, fence, or container that maintains certain species within an
ecological system.
endemic. A species that is found in one and only one particular geographic location.
epiphytes. Plants that live on other plants, but are not parasitic.
eutrophication. Eutrophication is extreme productivity from addition of nutri-
ents in a body of water, frequently due to runoff from the land, which causes
dense vegetation and death of animals from lack of oxygen.
fronds. The leaves of a fern.
greenhouse gas. A gas that contributes to the greenhouse effect by absorbing
infrared radiation, such as carbon dioxide.
greenhouse gas emissions. Releases of greenhouse gases into the atmosphere.
gross primary production. The amount of organic chemical energy as biomass
that primary producers create in a given length of time.
homeostasis. Maintain internal conditions within a range of acceptable extremes.
46 GLOSSARY
hydrologic cycle. The circulation of water throughout ecological systems and the
atmosphere through evaporation, transpiration, and precipitation.
hyperaccumulators. A plant that accumulates trace elements from its environ-
ment in concentrations higher than those found in the environment.
leaf area index. The ratio of total upper leaf surface area divided by land surface
area.
net primary production. The amount of gross primary production minus respira-
tion in a given length of time.
nitrogen cycle. The nitrogen cycle describes the circulation and chemical reac-
tions of nitrogen in ecological systems.
nitrogen fixation. Nitrogen fixation is the conversion of atmospheric nitrogen
(N2) to ammonia (NH3)
nutrient dynamics. The changes in nutrient concentrations in compartments
over time.
particulate matter. Living or nonliving material suspended in the water column.
phenological. Periodic biological phenomena that occur seasonally or annually.
photosynthesis. The process by which carbon dioxide and water are converted
into carbohydrates by green plants, algae, and certain bacteria using energy from
the sun.
photosynthetically active radiation. The amount of light available for photosyn-
thesis, in the 400 to 700 nanometer wavelength range.
pollutants. A human-produced or natural substance or condition that contami-
nates air, water, or soil.
positive feedback. Amplification of an effect by its own influence on the process
that gives rise to it.
proxy. A variable that stands in for another variable.
regeneration. The recycling and release of nutrients from organic matter by
decomposer organisms.
respiration. Cellular respiration is a process in organisms involving the trans-
duction of energy, typically with the intake of oxygen and the release of carbon
dioxide from the oxidation of complex organic substances.
seed bank. The storage of seeds, usually dormant, within the soils of most ecosystems.
seston. Minute living and non-living particulate matter suspended in the water
column.
tragedy of the commons. A phenomenon where individual users acting indepen-
dently and rationally in their own self-interest behave contrary to the common
good of all users by depleting or polluting a resource.
water column. Open water between the surface and the sediment.
xylem. Xylem is supporting and water-conducting tissue, which brings water and
nutrients from the roots to the rest of the plant.
zooplankton. Plankton, or drifting organisms, consisting of small animals and
the immature stages of larger animals.
Index
Absorbed photosynthetically active Free-Air Carbon dioxide Enrichment
radiation (APAR), 3031 (FACE), 2829, 31
Accumulators. See Hyperaccumulators
Acid precipitation, 24 Global carbon cycle, 27
Arsenic contamination Global climate change, 27, 32, 34,
in fronds, 2223 35, 3942
in roots, 2223 Global temperatures, 27, 32
in soils, 22 Global warming, 42
Arsenic toxicity, 23 Greenhouse gas emissions, 27
Assimilation, 1, 5, 8 Gross primary production (GPP), 29
Atmospheric carbon dioxide, 39 Gulf of Mexico dead zone, 14, 15
Biodiversity, 39, 42 Homeostasis

Biogeochemistry, 1213 in ecological systems, 7, 8, 13, 14,
Bormann, Herbert, 810, 14 16, 19, 27
in individual organisms, 19
of nutrients, 8
Carbon dioxide
Hydrologic cycle, 11
concentrations, 27
Hyperaccumulators, 21, 22
emissions, 24
increasing atmospheric, 2742
Inorganic materials, 11
Carbon sink, 27
Carbon source, 27
Leaf area index (LAI), 31
Climate
Leaf mortality, 36, 37
defined, 40
Leaf production, 36, 37
change, 27, 32, 34, 35, 39, 40
Likens, Gene, 89, 11, 14
Cloud water, 3639
Compartments in
Metal percentage in soils, 20
ecological systems, 1, 8
Metal pollution, 19
Contaminants, 19
Monteverde Cloud Forest Reserve
(MCFR), 36
Decomposition, 1, 6 Mutually agreed upon coercion, 26
Ecological mismatch, 35 Nadkarni, Nalini, 3538

Ecological systems National Climatic Data Center, 33
homeostasis of, 27 Net primary production (NPP),
filter wastes, 1926 2931
Epiphyte mats, 36, 3839 Nickel concentrations
Epiphytes in fruits, 21
mortality, 38 in plant tissues, 21, 23
viability or growth of, 36 in S. acuminate latex, 21
Eutrophication, 14 in soil, 21
48 INDEX
Nitrogen fixation, 1, 8 Seston, 3

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

FOR THE Christopher J. Paradise A. Malcolm Campbell

Christopher J. Paradise, PhD

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First published in 2016 by

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

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Printed in the United States of America

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

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

= loss from WC to wall

no fish low fish medium fish high fish

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

fraction of particulate phosphorus when fish were present than when

Figure 2 Simplified schematic of the phosphorus cycle. Inorganic

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

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

Figure 3 Annual particulate matter output in kilograms of dry mass

scientists further prevented regrowth of vegetation the following 3.5 years

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

elements in particulate matter were obtained by multiplying data from

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

Ethical, Legal, Social Implications: The Gulf of Mexico

Sabina Lake Lake Calcasieu

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

agricultural and fishing industries have vested economic interests at play,

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

Ecological Systems Can

In Chapter 1, it was shown how nutrients are exported from terrestrial

Table 1 Percentages of metals in soils from

Although metals may remain in ecological systems for hundreds of years,

and 0.48% in flowers. Psychotria douarrei, also from New Caledonia,

cells. Hyperaccumulation by plants might be an emergent property of eco-

in fronds quickly reached 6,000 ppm and ultimately climbed to over

concentrations. In this and other arsenic accumulators, arsenate (AsO43)

Ethical, Legal, Social Implications:

The tragedy is the dilemma that arises as individuals, acting indepen-

without a change in human values, human livelihoods, or ideas of moral-

Figure 6 The FACE experimental site at the Duke University Forest.

The scientists involved in this study work at several sites in forests in

Norby and his colleagues presented evidence that suggests a feedback

Nadkarni and Solano monitored leaves of plants from four representa-

Table 2 Results of greenhouse experiment on

Source: From Nadkarni and Solano, 2002.

Exposure to conditions with less cloud water, which is a prediction for

Ethical, Legal, Social Implications: There Is a

deviation from 30 year (19611990) mean temperature (C)

Figure 7 Anomalies for mean summer temperatures (July, August,

Biodiversity, 39, 42 Homeostasis

Ecological mismatch, 35 Nadkarni, Nalini, 3538

Nitrogen fixation, 1, 8 Seston, 3

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

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