Biodiversity: Tropical rainforests are areas of extremely high biodiversity

compared to other ecosystems. In the topical rainforests of Borneo, scientists

have documented more than 15,000 plant species, including 2,500 species of
orchids! Biologists estimate that tropical rainforests contain about 50% of the
worlds terrestrial plant and animal species, yet they encompass only about
6% of the worlds land area.

While tropical rainforests around the world have many similarities in their
climates and soil composition, each regional rainforest is unique. You will
not find precisely the same species living in all the tropical rainforests around
the world. For example, the species in African tropical rainforests are not the
same as the species living in the tropical rainforests of Central America.
However, the different species play similar roles within their specific regional
rainforest.

Population And Community Development And

Structure
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Smithsonian Tropical Research Institute (STRI)
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Tropical rainforests are distinguished not only by a remarkable richness of biota but
also by the complexity of the interrelationships of all the plant and animal inhabitants
that have been evolving together throughout many millions of years. As in all
ecosystems, but particularly in the complex tropical rainforest community, the
removal of one species threatens the survival of others with which it interacts. Some
interactions are mentioned below, but many have yet to be revealed.
General structure of the rainforest
Plants with similar stature and life-form can be grouped into categories
called synusiae, which make up distinct layers of vegetation. In tropical rainforests the
synusiae are more numerous than in other ecosystem types. They include not only
mechanically independent forms, whose stems are self-supporting, and saprophytic
plants but also mechanically dependent synusiae such as climbers, stranglers,
epiphytes, and parasitic plants. An unusual mix of trees of different sizes is found in
the tropical rainforest, and those trees form several canopies below the uppermost
layer, although they are not always recognizably separate layers. The upper canopy of
the tropical rainforest is typically greater than 40 metres above ground.
The tropical rainforest is structurally very complex. Its varied vegetation illustrates the
intense competition for light that goes on in this environment in which other climatic
factors are not limiting at any time of year and the vegetation is thus allowed to
achieve an unequaled luxuriance and biomass. The amount of sunlight filtering
through the many layers of foliage in a tropical rainforest is small; only about 1
percent of the light received at the top of the canopy reaches the ground. Most plants
depend on light for their energy requirements, converting it into chemical energy in
the form of carbohydratesby the process of photosynthesis in their chlorophyll-
containing green tissues. Few plants can persist in the gloomy environment at ground
level, and the surface is marked by a layer of rapidly decomposing dead leaves rather
than of small herbaceous plants. Mosses grow on tree butts, and there are a few forbs
such as ferns and gingers, but generally the ground is bare of living plants, and even
 shrubs are rare. However, tree seedlings and saplings are abundant; their straight
stems reach toward the light but receive too little energy to grow tall enough before
food reserves from their seeds are exhausted. Their chance to grow into maturity
comes only if overhanging vegetation is at least partially removed through tree death
or damage by wind. Such an occurrence permits more solar radiation to reach their
level and initiates rapid growth and competition between saplings as to which will
become a part of the well-lit canopy.

Vegetation profile of a tropical rainforest.
Encyclopdia Britannica, Inc.

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Gaps in the canopy of a tropical rainforest provide temporarily well-illuminated
places at ground level and are vital to the regeneration of most of the
forests constituent plants. Few plants in the forest can successfully regenerate in the
deep shade of an unbroken canopy; many tree species are represented there only as a
 population of slender, slow-growing seedlings or saplings that have no chance of
growing to the well-lit canopy unless a gap forms. Other species are present, invisibly,
as dormant seeds in the soil. When a gap is created, seedlings and saplings accelerate
their growth in the increased light and are joined by new seedlings sprouting from
seeds stored in the soil that have been stimulated to germinate by light or by
temperature fluctuations resulting from the suns shining directly on the soil surface.
Other seeds arrive by various seed-dispersal processes (see below). A thicket of
regrowth rapidly develops, with the fastest-growing shrubs and trees quickly shading
out opportunistic, light-demanding, low-growing herbaceous plants and becoming
festooned with lianas. Through it all slower-growing, more shade-tolerant but longer-
lived trees eventually emerge and restore the full forest canopy. The trees that initially
fill in the gap in the canopy live approximately one century, whereas the slower-
growing trees that ultimately replace them may live for 200 to 500 years or, in
extreme cases, even longer. Detailed mapping of the trees in a tropical rainforest can
reveal the locations of previous gaps through identification of clumps of the quicker-
growing, more light-demanding species, which have yet to be replaced by trees in the
final stage of successional recovery. Local, natural disturbances of this sort are vital to
the maintenance of the full biotic diversity of the tropical rainforest (see Sidebar:
Rainforest Regeneration in Panama).
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Just as tropical rainforest plants compete intensely for light above ground, below
ground they vie for mineral nutrients. The process of decomposition of dead materials
is of crucial importance to the continued health of the forest because plants depend on
rapid recycling of mineral nutrients. Bacteriaand fungi are primarily responsible for
this process. Some saprophytic flowering plants that occur in tropical rainforests rely
on decomposing material for their energy requirements and in the process use and
later release minerals. Some animals are important in the decomposition process; for
example, in Malaysia termites have been shown to be responsible for the
decomposition of as much as 16 percent of all litter, particularly wood. Most trees in
the tropical rainforest form symbiotic mycorrhizal associations with fungi that grow
in intimate contact with their roots; the fungi obtain energy from the tree and in turn
provide the tree with phosphorus and other nutrients, which they absorb from the soil
very efficiently. A mat of plant roots explores the humus beneath the rapidly
decomposing surface layer of dead leaves and twigs, and even rotting logs are invaded
by roots from below. Because nutrients are typically scarce at depth but, along with
moisture, are readily available in surface layers, few roots penetrate very deeply into
the soil. This shallow rooting pattern increases the likelihood of tree falls during
storms, despite the support that many trees receive from flangelike plank buttresses
growing radially outward from their trunk bases. When large trees fall, they may take
 with them other trees against which they collapse or to which they are tied by a web
of lianas and thereby create gaps in the canopy.

An earthstar (Geastrum) puffball, growing on moist soil among mosses.
Larry WestThe National Audubon Society/Photo Researchers

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Tree growth requires substantial energy investment in trunk development, which some
plants avoid by depending on the stems of other plants for support. Perhaps the most
obvious adaptation of this sort is seen in plants that climb from the ground to the
uppermost canopy along other plants by using devices that resemble grapnel-like
 hooks. Lianas are climbers that are abundant and diverse in tropical rainforests; they
are massive woody plants whose mature stems often loop through hundreds of metres
of forest, sending shoots into new tree crowns as successive supporting trees die and
decay. Climbing palms or rattans (Calamus) are prominent lianas in Asian rainforests,
where the stems, which are used to make cane furniture, provide a valuable economic
resource.

Lianas in a tropical rainforest. The vascular tissues of lianas are modified primarily for water
Gary Braasch

Epiphytes are particularly diverse and include large plants such as orchids, aroids,
bromeliads, and fernsin addition to smaller plants such as algae, mosses, and lichens.
In tropical rainforests epiphytes are often so abundant that their weight fells trees.
Epiphytes that grow near the upper canopy of the forest have access to bright sunlight
but must survive without root contact with the soil. They depend on rain washing over
them to provide water and mineral nutrients. During periods of drought, epiphytes
undergo stress as water stored within their tissues becomes depleted. The diversity of
epiphytes in tropical deciduous forests is much less than that of tropical rainforests
because of the annual dry season (see Sidebar: Life in a Bromeliad Pool).

Epiphyte bromeliads (Vriesea).
Luiz Claudio Marigo/Bruce Coleman Ltd.

TEST YOUR KNOWLEDGE

Animal Adventures: Fact or Fiction?

Parasitic flowering plants also occur. Hemiparasitic mistletoes attached to tree

branches extract water and minerals from their hosts but carry out their
own photosynthesis. Plants that are completely parasitic also are found in tropical
rainforests. Rafflesia, in Southeast Asia, parasitizes the roots of certain lianas and
produces no aboveground parts until it flowers; its large orange and yellow blooms,
nearly one metre in diameter, are the largest flowers of any plant.
 CONNECT WITH BRITANNICA

Stranglers make up a type of synusia virtually restricted to tropical rainforests. In this

group are strangler figs (Ficus), which begin life as epiphytes, growing from seeds left
on high tree branches by birds or fruit bats. As they grow, they develop long roots that
descend along the trunk of the host tree, eventually reaching the ground and entering
the soil. Several roots usually do this, and they become grafted together as they
crisscross each other to form a lattice, ultimately creating a nearly complete sheath
around the trunk. The host trees canopy becomes shaded by the thick fig foliage, its
trunk constricted by the surrounding root sheath and its own root system forced to
compete with that of the strangling fig. The host tree is also much older than the
strangler and eventually dies and rots away, leaving a giant fig tree whose apparent
 trunk is actually a cylinder of roots, full of large hollows that provide shelter and
breeding sites for bats, birds, and other animals (see Sidebar: Apartments of the
Rainforest). Stranglers may also develop roots from their branches, which, when they
touch the ground, grow into the soil, thicken, and become additional trunks. In this
way stranglers grow outward to become large patches of fig forest that consist of a
single plant with many interconnected trunks.

Relationships between the flora and fauna

Some of the tallest trees and lianas, and the epiphytes they support, bear flowers and
fruits at the top of the rainforest canopy, where the air moves unfettered by vegetation.
They are able to depend on the wind for dispersal of pollen from flower to flower, as
well as for the spreading of fruits and seeds away from the immediate environment of
the parent plant (see Sidebar: Flying Trees). Ferns, mosses, and other lower plants
also exploit the wind to carry their minute spores. However, a great many flowering
plants, including many that grow in the nearly windless environment of the
understory, depend on animals to perform these functions. They are as dependent on
animals for reproductive success as the animals are on them for foodone example of
the mutual dependence between plants and animals (see Sidebar: No Rainforest, No
Brazil Nuts).
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Many rainforest trees have sizable seeds from which large seedlings emerge and thrust
their way through the thick mat of dead leaves on the dark forest floor. They develop
tall stems, using food reserves in the seed without having to rely on sunlight, which is
usually too dim, to meet their energy requirements. Because large seeds cannot be
dispersed by the wind, these plants depend on a variety of animals to perform this
function and have evolved many adaptations to encourage them to do so. Fruit
bats are attracted by fragrant, sweet fruits typically borne conspicuously and
conveniently on the outer parts of the tree canopy; the mango (Mangifera indica),
native to the rainforests of India, provides a good example. The bats not only feed on
fruits as they hang from the trees but also may carry a fruitaway to another perch,
where they eat the flesh and drop the seed. Smaller fruits may be swallowed whole,
the seeds passing through the gut intact and being voided at a distance. The ground
 beneath trees used by fruit bats as a roost is commonly thick with seedlings of fleshy,
fruit-bearing trees (seeSidebar: Hitching a Ride).

Mango (Mangifera indica).

Robert C. HermesThe National Audubon Society Collection/Photo Researchers

A variety of birds eat fleshy fruits also, voiding or regurgitating the unharmed seeds.
Birds of different sizes are typically attracted to similarly scaled fruits, which are
carried on stems of appropriate thickness and strength. For example, large pigeons in
New Guinea feed preferentially on larger fruits borne on thicker stems that can bear
not only the weight of the fruit but also the weight of the large bird; smaller pigeons
tend to feed on smaller fruits borne on thinner twigs. In such a manner, the diverse
plant community is matched by a similarly diverse animal community in
interdependence.
Terrestrial mammals also help to disperse seeds. In many cases this has favoured the
positioning of flowers and fruits beneath the canopy on the trunks of trees accessible
to animals unable to climb or fly, an adaptation called cauliflory. In some cases fruits
are grown in the canopy but drop as they ripen, opening only after they fall to attract
ground-dwelling animals that will carry them away from the parent tree.
The durian fruit Durio zibethinus of Southeast Asian rainforests is an example; its
fruits are eaten and its seeds dispersed by a range of mammals, including pigs,
elephants, and even tigers.
Many other animals, from ants to apes, are involved in seed dispersal. In the Amazon
basin of Brazil, where large areas of tropical rainforest are seasonally flooded, many
trees produce fruit attractive to fish, which swallow them whole and void the seeds
(see Sidebar: Vegetarian Piranhas). Squirrels are also important seed dispersers in
parts of South America. In the tropical rainforests of northeastern
Australia, cassowaries are responsible for generating mixed clumps of tree seedlings
of several species that grow from their dung sites.
It is important for seeds to be spread away from parent plants, both to allow seedlings
to escape competition with the parent and to expand the range of the species. Another
capacity important to seed survival, particularly in the diverse tropical rainforest
community, involves the evasion of seed predators. Many different beetles and other
insects are specialized to feed on particular types of seed. Seeds concentrated beneath
a parent plant are easy for seed predators to locate. Seeds that are carried away to
areas occupied by different plant speciesand different seed predatorsare more
likely to survive (see Sidebar: Hitching a Ride).
In addition to dispersing seeds, animals are vital to tropical rainforest reproduction
through flower pollination. Many insects such as bees, moths, flies, and beetles as
well as birds and bats carry out this activity. Birds such as
the hummingbirds of South and Central America and the flower-peckers of Asia have
adaptations that allow them to sip nectar from flowers. In the process they
inadvertently become dusted with pollen, which they subsequently transport to other
flowers, pollinating them. The plants involved also show special adaptations in flower
structure and colour. Most flowers pollinated by birds are red, a colour highly visible
to these animals, whereas flowers pollinated by night-flying moths are white or pink,
and those pollinated by insects that fly during the day are often yellow or
orange. Bats are important pollinators of certain pale, fragrant flowers that open in the
evening in Asian rainforests (see also Sidebar: Bat-Loving Flowers).

Biological Productivity
Of all vegetation types, tropical rainforests grow in climatic conditions that are least
limiting to plant growth. It is to be expected that the growth and productivity (total
amount of organic matter produced per unit area per unit time) of tropical rainforests
would be higher than that of other vegetation, provided that other factors such as soil
fertility or consumption by herbivorous animals are not extremely low or high.
Various methods are employed to assess productivity. Gross primary productivity is
the amount of carbon fixed during photosynthesis by all producers in the ecosystem.
However, a large part of the harnessed energy is used up by the metabolic processes
of the producers (respiration). The amount of fixed carbon not used by plants is
called net primary productivity, and it is this remainder that is available to various
consumers in the ecosysteme.g., the herbivores, decomposers, and carnivores. Of
course, in any stable ecosystem there is neither an accumulation nor a diminution in
the total amount of organic matter present, so that overall there is a balance between
the gross primary productivity and the total consumption. The amount of organic
matter in the system at any point in time, the total mass of all the organisms present, is
called the biomass. (For further discussion of productivity, see biosphere: Resources
of the biosphere.)
The biomass of tropical rainforests is larger than that of other vegetation. It is not an
easy quantity to measure, involving the destructive sampling of all the plants in an
area (including their underground parts), with estimates made of the mass of other
organisms belonging to the ecosystem such as animals. Measurements show that
tropical rainforests typically have biomass values on the order of 400 to 700 metric
tons per hectare, greater than most temperate forests and substantially more than other
vegetation with fewer or no trees. A measurement of biomass in a tropical deciduous
forest in Thailand yielded a value of about 340 metric tons per hectare.

Increase in biomass over the period of a year at one rainforest site in Malaysia was
estimated at 7 metric tons per hectare, while total litter fall was 14 metric tons,
estimated mass of sloughed roots was 4 metric tons, and total live plant matter eaten
by herbivorous animals (both invertebrate and vertebrate) was about 5 metric tons per
hectare per year. These values add up to a total net production of 30 metric tons per
hectare per year. Respiration by the vegetation itself was estimated at 50 metric tons,
so that gross primary productivity was about 80 metric tons per hectare per year.
Compared with temperate forests, these values are approximately 2.5 times higher for
net productivity and 4 times higher for gross productivity, the difference being that the
respiration rate at the tropical site was 5 times that of temperate forests.
Despite the overall high rates of productivity and biomass in tropical rainforests, the
growth rates of their timber trees are not unusually fast; in fact, some temperate trees
and many smaller herbaceous plants grow more rapidly. The high productivity of
tropical rainforests instead results in their high biomass and year-round growth. They
also have particularly high levels of consumption by herbivores (see Sidebar: Eating
the Rainforest), litter production, and especially plant respiration.

Plants Versus Animals: Do They Deal with

Stress in Different Ways?1
Raymond B. Huey Margen Carlson Lisa Crozier Melanie Frazier Hayden HamiltonChristopher
Harley Anhthu Hoang Joel G. Kingsolver

Integrative and Comparative Biology, Volume 42, Issue 3, 1 July 2002, Pages 415
423,https://doi.org/10.1093/icb/42.3.415

Published:

01 July 2002


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Abstract
Both plants and animals respond to stress by using adaptations that help them evade, tolerate, or
recover from stress. In a synthetic paper A. D. Bradshaw (1972) noted that basic biological
differences between plants and animals will have diverse evolutionary consequences, including those
influencing how they deal with stress. For instance, Bradshaw argued that animals, because they have
relatively well-developed sensory and locomotor capacities, can often use behavior and movement to
evade or ameliorate environmental stresses. In contrast, he predicted that plants will have to
emphasize increased physiological tolerance or phenotypic plasticity, and also that plants should
suffer stronger selection and show more marked differentiation along environmental gradients. Here
we briefly review the importance of behavior in mitigating stress, the behavioral capacities of
animals and plants, and examples of plant responses that are functionally similar to behaviors of
animals. Next, we try to test some of Bradshaw's predictions. Unfortunately, critical data often
proved non-comparable: plant and animal biologists often study different stressors
(e.g., water versus heat) and measure different traits (photosynthesis versus locomotion).
Nevertheless, we were able to test some of Bradshaw's predictions and some related ones of our own.
As Bradshaw predicted, the phenology of plants is more responsive to climate shifts than is that of
animals and the micro-distributions of non-mobile, intertidal invertebrates (plant equivalents) are
more sensitive to temperature than are those of mobile invertebrates. However, mortality selection is
actually weaker for plants than for animals. We hope that our review not only redraws attention to
some fascinating issues Bradshaw raised, but also encourages additional tests of his predictions. Such
tests should be informative.
Issue Section:

Regular Article

INTRODUCTION

Perhaps then zoologists might look at plants, and botanists look at animals, and we may find that
evolution in some animals is like that in plants, and evolution in some plants like that in animals,
particularly when their properties overlap.

A. D. Bradshaw (1972, p. 25)

Environmental changes influence the performance and fitness of organisms. Some environmental
changes are beneficial, but most will be stressful (Fisher, 1958). To ameliorate stressful changes,
organisms can adopt three general and non-exclusive mechanisms (Larcher et al., 1973, p. 231).
First, they can evade or reduce the stress by using behavior (e.g., changing habitats or temporal
activity patterns) or dormancy. Second, they can evolve greater resistance to stress (via increased
tolerance, greater plasticity, reduced sensitivity). Third, if actually damaged, they can
activate recovery mechanisms (e.g., regeneration of damaged tissues, cellular stress responses).

Animals and plants use all three strategies, but are thought to differ in which of these strategies
they emphasize. In particular, animals often rely on behavior as their first line of defense against
environmental challenges (Bartholomew, 1964). Such behavioral adjustments can sometimes
completely mitigate a challenge, thus obviating the need to activate alternative anti-stress
responses. However, plants cannot run away from a stress and are thus more likely to emphasize
dormancy, stress-resistance, or stress-recovery mechanisms (see below).

In 1972 A. D. Bradshaw published a seminal paper Some of the evolutionary consequences of

being a plant. Noting that plants are relatively immobile and have fewer behavioral options than
do animals, Bradshaw proposed that plants should tolerate a broader range of environmental
conditions, show greater phenotypic plasticity, and experience stronger selection in nature.
Subsequently, other botanists have extended Bradshaw's theme (Harper, 1977; Levin,
1978; Niklas, 1992). Moreover, as Bradshaw noted (1972, p. 26), some of his predictions can be
readily applied to comparisons involving sessile (i.e.,plant-like) versus mobile animals.

Here we have three general goals. First, we briefly review how and why behavior is a useful way
to deal with environmental stress. Second, we evaluate whether plants are really as immobile as
sometimes assumed. Third, we review some of Bradshaw's predictions and try to evaluate their
validity.

We made some progress towards the first two goals, but have had limited success with the third.
Quantitative tests of Bradshaw's predictions proved difficult to implement, not merely because
the relevant literatures are huge and scattered, but mainly because the studies often proved non-
comparable.The problem here is simply that plant and animal biologists often measure different
traits and different environmental variables. Nevertheless, the issues raised by Bradshaw
(1972) are fascinating, relevant, and deserving of additional testing. We hope that our paper will
acquaint animal biologists with Bradshaw's paper. Although frequently cited by botanists, it has
not (to our knowledge) been cited by a zoologist.

ON THE UTILITY OF BEHAVIOR

Regulatory behavior (by which we mean the capacities to sense the environment and to respond
with movement) can be an effective way of dealing with changing environments. For example,
such behavior can help an organism control its physiological vulnerability to abiotic factors by
reducing the odds that it encounters stressful or lethal conditions (Bartholomew,
1964; Stevenson, 1985) and by increasing the proportion of time that it spends at physiologically
optimal conditions (Huey et al., 1989).

Behavioral evasion can also influence the outcome of biotic interactions (Helfman, 1986).
Consider a frog being stalked by a heron. The frog can use its senses to detect the predator at a
distance and then take evasive action. In contrast, consider a pumpkin that is in the path of a
migrating elephant. The inevitable happens: the pumpkin is transmuted into a squash.

The ability to detect and evade biotic and abiotic threats is oftenbut not alwaysadaptive on
balance. Movement expends energy and can attract predators (Roth, 1986), and increased
movement capacity may evolve only at the sacrifice of reproductive effort (Chai and Srygley,
1990). In any case, high mobility is not inevitably the sine qua non of evolution. Sloths and
barnacles are derived from mobile ancestors, implying that selection sometimes favors reduced
mobility of animals (i.e., becoming more plant like).

ARE PLANTS STATIONARY?

I shall publish in a month or two a book on the Movements of Plants. I will send you a copy,
but I fear it is much too special for anyone but a physiological botanist to care about. I have long
thought that old men, like myself, ought to write only on confined & easy subjects.

Charles Darwin, Oct. 7, 1888

(unpublished letter to Ernst Krause)

Bradshaw (1972, p. 25) noted that some plants are animal like and that some animals are plant
like. Nevertheless, he (1972, p. 26) generalized that animals are mobile and that plants are
stationary: So a plant cannot move from one environment into another in the course of its
lifetime, in the way that an animal may traverse numerous different environments in the course
of a day. This generalization is obviously valid, but what strikes us (as zoologists looking at
plants) is that some plant responses do involve physical movement and are remarkably animal-
like in effect, though not in scale.
Let us continue with the theme of behavioral thermoregulation. Stevenson (1985) showed that
many animals gain considerable control over body temperature by using movement to select
habitats that differ in operative temperatures (Huey, 1991), to restrict activity to times with
suitable temperatures (Porter and Tracy, 1983), or to orient to heat sources and sinks (Heath,
1965). (Nevertheless, not all animals that have such regulatory capacities always use them in
nature [Feder et al., 1996; but see Jones et al., 1987].)

What about plants? Are they passive to changes in the thermal environment? In fact, some plants
have responses that achieve much the same ends as the overt locomotor adjustments of animals.
As Bazzaz (1991) noted, some plants can effectively select habitats by growing towards some
desirable resource or away from an environmental stressor. Some plants use the ratio of red:far-
red wavelengths of light, which depends on the degree of shading, as a cue of the presence of
other plants in the immediate environment; and they can adjust their growth accordingly and
adaptively (Dudley and Schmitt, 1995). Neotropical stilt palms (Socratea exorrhiza) are reported
(Leopold et al., 2000) to move on their stilts towards light gaps: alas, this is apparently a tropical
myth (A. Henderson, personal communication).

Climbing and clonal plants can literally crawl across the environment, selecting habitats much
like animals. In fact, such movements are sometimes called foraging (Bazzaz, 1991; Harper,
1986; Ray, 1992). Remarkably, connected ramets living in different habitats can even share
nutrients and water (Pitekla and Ashmun, 1985; Salzman, 1985).

To some extent, plants may target dispersal of progeny toward favorable habitats (Bazzaz, 1991).
Of course, fruits and seeds of many plants are adapted for dispersal by animals: in most cases,
such dispersal is undoubtedly haphazard. However, seed dispersal by some ants (Kalisz et al.,
1999), and perhaps also mistletoe seeds by birds, is not.

Plants can orient their leaves and flowers towards or away from the sun (Stanton and Galen,
1989), in ways functionally similar the orientation movements of animals (Heath,
1965; Stevenson, 1985). Ehleringer and Forseth (1980) showed that the leaves of many plants
change their orientation during the day, maintaining a perpendicular orientation to the sun's rays,
and thereby enhance photosynthesis (rather than modulate body temperature as in lizards).
However, water stressed plants reposition their leaves to a vertical orientation (Ehleringer and
Forseth, 1980), reducing heat gain and thus conserving water. Solar tracking by flowers of alpine
buttercups (Ranunculus) provides a heat reward to insect pollinators as well as to increase seed
set (Stanton and Galen, 1989).

Charles Darwin (see quote above) and his son Francis wrote a book on movement of plants
(Darwin and Darwin, 1880) and included a chapter on sleeping in plants. The leaves of many
plants droop at night, and this had long been interpreted as a way of conserving energy. Darwin
suspected that such sleeping postures actually had a thermoregulatory function: a leaf held
horizontal and exposed to the clear night sky would lose heat by radiation and potentially suffer
frost damage. To test this hypothesis, the Darwins put out plants at night, allowing some leaves
to sleep, but holding others horizontal. Leaves allowed to sleep had much lower frost damage
than did leaves held horizontal. In Oxalis actosella, for example, 92% of the leaves that were
sleepers survived a cold night, whereas only 37% of the non-sleepers survived (P = 0.003,
by a Fisher exact test, of course). Although competing explanations for the phenomena have
been offered (Bunning, 1967), our point is that leaves can move, in ways that are functionally
similar to the orientation movements of many animals.

Some plants even shift the solar orientation of chloroplasts within their leaves, even when the
leaves do not move. Duckweed is a small-leaf plant that is aquatic, with the leaves held
horizontally on the water surface. The chloroplasts are oriented perpendicularly to the light in
dim light, thereby maximizing uptake of light; but in bright light, which is potentially damaging,
the chloroplasts rotate so that they are parallel to the light, thereby minimizing light exposure.
The sensory mechanism underlying such remarkable chloroplast movement has recently been
described in Arabidopsis(Kagawa et al., 2001).

Finally, even though they themselves are relatively sessile, plants can take advantage of an
animal's mobility. Plants of course routinely manipulate animal movement for pollination and
dispersal, but some plants even manipulate animal movement for self-defense (Kessler and
Baldwin, 2001). When attacked by herbivorous spider mites, lima beans secrete volatile
compounds that attract predatory mites, which then attack the herbivorous mites, thus providing
an indirect defense for the plant (Dicke, 1999). Further, some plants (sundews, Venus flytrap,
pitcher plants) trap mobile animals (Hart, 1990).
We have highlighted only a few examples of plant movements (Hart, 1990). We are not claiming
that plants can match the sensory and locomotor capacities of animals, and we wish to avoid the
semantic debate of whether plant movements should be called behavior. What does matter here
is that some plants movements achieve many of the same functional ends as those of animals.
Plants may not move as far or as quickly as animals, but some are hardly immobile. Mobility
differences are often a matter of scale.

DO PLANTS AND ANIMALS DEAL WITH STRESS IN

DIFFERENT WAYS?

Given that plants are relatively stationary and thus forced to experience environmental
pressures, Bradshaw (1972) made several predictions as to ways plants and animals should differ
in their ecology and evolution. We focus on the subset of his predictions relating to
environmental tolerance, and we've added a few predictions that are philosophically consistent
with his perspective.

Prediction: plants have broader tolerance ranges than

do animals

Frequent exposure to extreme environments may select for enhanced tolerance capacity
(Hoffmann and Parsons, 1991). Accordingly, plants should tolerate broader ranges of
environmental conditions than would animals, simply because plants won't be able to evade a
stressor.

This prediction is testable in two ways. First, the premise that plants experience wider ranges of
conditions could be tested by determining whether, for example, plants have actually experience
more variable body temperatures over time than do sympatric ectothermal animals.
Unfortunately, such data have never been gathered systematically at any locality. Doing so will
be difficult because some individual plants will show much greater regional heterothermy
(i.e., the temperature differences between different parts of the plant) than do animals (see fig.
4.7 in Krner, 1999), making it relatively hard to characterize the temperature of a plant.
Second, data on actual tolerance ranges (i.e., the range between upper and lower lethal
temperatures) of plants and of animals could be compared directly, ideally for organisms studied
at the same sites. Implementing this test proved frustrating. Much of the literature on vertebrate
ectotherms measures critical thermal limits (temperatures at which the righting response is
lost), which have no direct equivalents in the plant literature. Lethal temperatures are commonly
measured in both groups, but would have to be standardized for exposure time and acclimation
condition. Supercooling points are also measured in some plants and animals (Lee et al.,
1995; Levitt, 1980), but often in quite different ways (R. E. Lee, personal communication). So
one would have to validate (study by study) that the techniques and end-points being used were
comparable. Moreover, any meaningful analysis would need to compare tolerance temperatures
of multiple stages of plants versusanimals, as tolerance can be strongly stage dependent, on
species collected from the same area or habitat.

Given these and other complications, we made only a cursory search of the literature. Consistent
with Bradshaw's reasoning, some plants (especially seeds) are extraordinarily cold tolerant (see
table 3.1 in Levitt, 1972) relative to insects (table 5.7 in Withers, 1992) and reptiles (table 1
in Heatwole, 1976). Similarly, some plants (especially seeds) are extraordinarily heat tolerant
(table 11.2 in Levitt, 1972) even relative to desert reptiles (Heatwole, 1976). However, whether
plants generally have broader tolerance ranges than do sympatric animals remains to be
quantified systematically.

Prediction: plants should emphasize biochemical

adaptations more than do animals

Bradshaw (1972) did not discuss specific physiological mechanisms. However, if plants have
evolved greater physiological tolerance than animals (above), then their enhanced tolerance
should be evident at the biochemical level.

Organisms that experience a wide range of body temperatures may possess eurythermal
enzymes, which operate efficiently over a wide range of temperatures (Somero, 1995). An
enzyme's thermotolerance is often assessed by measuring the thermal sensitivity of its Km value
(Patterson and Graham, 1987; Somero et al., 1996; Teeri, 1980), which is the concentration of
substrate needed for an enzyme to operate at half maximal velocity. Unfortunately, we know of
no direct comparisons of the thermal sensitivity of Km values of plants versus animals.

An organism's thermal tolerance could also be increased by evolving multiple variants of a given
enzyme, or isozymes, each with a different thermal optimum (Baldwin and Hochachka,
1970; Hochachka and Somero, 1984; Lin and Somero, 1995). Few animals use isozymes,
perhaps because of the genetic constraints of diploidy (Hochachka and Somero, 1984; Somero,
1995). Interestingly, many plants are polyploids (Ramsey and Schemske, 1998) and might
therefore be more likely to evolve thermally distinct isozymes. However, whether plants
commonly use isozymes to cope with wide thermal ranges is unclear (Lutova, 1995; Teeri,
1980), though some plants have isozymes that fluctuate in quantity with seasonal changes
(Hausladen and Alscher, 1994).

On occasion organisms (mobile or not) are exposed to stressful or damaging temperatures, and
plants and animals synthesize special proteins that stabilize cells at such temperatures (Feder and
Hofmann, 1999). These proteins have quite diverse functions (e.g., stabilization of protein
structure, alteration of membrane fluidity, and prevention of cellular ice formation). Many such
proteins, particularly heat shock proteins, have been well studied in plants and animals
(Boston et al., 1996; Feder, 1999; Feder and Hofmann, 1999; Never et al., 1996; Patterson and
Graham, 1987; Sabehat et al., 1998; Somero, 1995). However, whether plants have relatively
heightened stress responses (e.g., faster or stronger synthesis of stabilizing proteins;
see Coleman et al., 1995) has not yet been determined. Suggestively, however, the temperature
that induces the synthesis of heat shock proteins (Buckley et al., 2001; Roberts et al.,
1997; Tomanek and Somero, 1999) is lower for intertidal mussels (sessile) than that for
sympatric snails (mobile) held under similar conditions.

Prediction: plants are more plastic than are animals

Organisms can adjust to environmental variation by altering their phenotypes (Schlichting and
Pigliucci, 1998), a phenomenon that is called phenotypic plasticity (or acclimation, when
reversible). Plasticity is often assumed to be an adaptation, though some studies are contradictory
(Huey et al., 1999; Leroi et al., 1994; Sultan and Bazzaz, 1993). In a classic paper, Bradshaw
(1965) called attention to the general importance of plasticity. Then in his 1972 paper, he
predicted that plants should have evolved relatively high levels of plasticity. In addition, the
modular structure and less targeted vegetative morphology of plants enable them to have much
greater morphological plasticity than could typical animals (Bazzaz, 1991; Sultan, 1987).

To test Bradshaw's (1972) prediction, we decided to use a useful index of Levins (1969) to
quantify plasticity (see also Kingsolver and Huey, 1998). For example, the slope of the
regression of lethal temperature on acclimation temperature quantifies the plasticity of the
thermal acclimation response. However, when we tried to compile data on plasticity from
literature, we ran into problems. First, and most importantly, plant and animal biologists often
study different environmental variables. This became conspicuous when we conducted a citation
search (Table 1) on phenotypic plasticity plus (plant or animal) plus one environmental
variable (temperature, water, soil, or light): the vast majority of the animal studies dealt with
temperature, whereas plant studies more commonly dealt with water, nutrients, or light (Sultan,
2001). Second, plant and animal biologists often measure very different traits
(e.g.,photosynthesis versus running speed). Third, animal biologists typically acclimate animals
to fixed temperature regimes, whereas plant biologists usually use cycling acclimation regimes.
This may trace to Went's (1957)classic studies, showing that plants grow better on cycling
regimes.

Despite these technical concerns, comparable data surely exist in the literature, but finding and
compiling these data will require a cautious search. So a rigorous test of Bradshaw's plasticity
hypothesis remains an open opportunity.

Prediction: plants show stronger inducible defenses

than do animals

Inducible defenses are a special case of phenotypic plasticity made directly in response to cues
from biotic agents such as predators, herbivores, or predators (Tollrian and Harvell, 1999); and
they are assumed to provide at least partial protection following attack. Bradshaw (1972) did not
specifically discuss inducible defenses, but subsequent workers (Harvell, 1990) have proposed
that inducible defenses might be especially favored in sessile and in clonal (or modular)
organisms such as plants or colonial invertebrates, which are unable to run away from predators.
Although many of the initial examples of inducible defenses came from studies of sessile
organisms (Harvell, 1990), inducible defenses are increasingly detected in mobile and in non-
clonal organisms (Relyea, 2001; Tollrian and Harvell, 1999). Moreover, immune responses,
which are a class of inducible defenses (Frost, 1999), are well developed in vertebrates, which
are neither sessile nor clonal. In any case, it is not obvious how to compare the magnitude of
different kinds of inducible defense.

Prediction: selection should be stronger on plants

than on animals

Because plants don't move through several habitats during their lifetimes but instead are
chronically exposed to a single one, they experience the world as more coarse-grained
(sensuLevins, 1968) than do animals. As a result, Bradshaw (1972, p. 26) predicted that selection
should be relatively strong on plants. Indeed, many of the classic examples of strong selection in
nature involve plants (Bradshaw, 1972).

To quantify the force of directional selection on a quantitative trait, we use the standardized
selection gradient (Lande and Arnold, 1983), which is the partial regression coefficient (linear)
of relative fitness on a trait score (in units of standard deviations). Kingsolver et al. (2001) have
recently compiled estimates of these coefficients
(http://www.bio.unc.edu/faculty/kingsolver/lab/).

Analyzing this data set, we partitioned s (absolute values) for morphological traits of plants,
vertebrates, versus invertebrates; and we've done separate analysis for studies that
measured mortality (viability) selection from those that measured non-mortality aspects of
selection (e.g., fecundity, fertility, or mating success).

Contrary to Bradshaw's prediction, plants do not in fact show stronger non-mortality selection
gradients on morphological traits than do animals (Savage test, P = 0.29). In fact, no trend in this
direction is even evident (Fig. 1a). Note, however, that plants show much weaker mortality
selection gradients than do animals (P < 0.001, Fig. 1b).
Why do the data contradict Bradshaw's prediction? Perhaps the plant data are somewhat
misrepresentative. Mortality selection on plants is probably strongest at the seed and seedling
stages (D. Schemske, personal communication), yet most plant studies to date are on adult
plants. For example, none of the available estimates of mortality selection involves seed
mortality, and at least 2/3 of the estimates involve mortality selection after establishment.

SPATIAL DISTRIBUTIONS

When confronted by a stress, a mobile organism can seek refuge in physically benign
microhabitats or abandon the area entirely. Plants and sessile animals do not enjoy this luxury
(though some use dormancy or seed banks) and may suffer widespread mortality from a single
stressful event (Bowers, 1982; Tsuchiya, 1983). All else being equal, sessile species may thus
have relatively restricted distributions along a stress gradient. Intertidal organisms provide a test
of this expectation.

Intertidal organisms face severe stresses (heat, desiccation) when exposed to the air during low
tide (Helmuth, 1998 and references therein). The highest tolerable position in the intertidal is
largely determined by an organism's ability to survive prolonged periods of emersion (Newell,
1979; Stillman and Somero, 1996). The upper limits of intertidal organisms vary predictably
with between-site patterns of physical stress (Leigh et al., 1987; Lewis, 1986; Wethey, 1984).
For example, on foggy coasts with heavy wave action, intertidal species can survive above the
actual limit of tidal excursion. However, on sunny, wave-protected shores, they survive only at
lower shore levels where aerial exposure is minimal (see also Lewis, 1964).

The inverse relationship between upper distributional limits and magnitude of physical stress
(e.g., substrate temperature) should be relatively strong for sessile intertidal species. In contrast,
upper limits of mobile species should be weakly related to between-site patterns of substrate
temperature as these species can retreat down-shore or into crevices.

To test this prediction, one of us (Harley, 2001) contrasted the upper intertidal limits of several
species of algae, sessile invertebrates, and mobile invertebrates at six localities ranging from the
foggy, wave exposed outer coast of Washington to the sunny, protected shores of Puget Sound.
Maximum rock temperatures in the mid-intertidal range from 26C on the outer coast to over
41C in Puget Sound. Consistent with expectation, upper limits are strongly negatively related to
maximum rock temperatures for algae and sessile invertebrates, but only weakly related to
maximum temperatures for mobile species. (Fig. 2). The upper distributional limits of sessile
species were significantly more sensitive to physical stress than were those of the mobile species
(Table 2).

PATTERNS THROUGH TIME

The phenology of annual events can change in response to shifts in climate. If plants are more
plastic than animals, then plant phenology should be relatively sensitive to inter-annual
variability in climate.

We test this prediction by re-analyzing a remarkable, multi-century dataset of phonologies

(e.g., time of bud burst for plants, of nest building for birds) recorded in southeastern England
(Sparks and Carey, 1995). Two patterns are evident. First, the timing of phenological events of
plants was much more closely tied to between-year climatic variability than was that of animals
(Fig. 3a, P 0.001, compiled from table 4 in Sparks and Carey, 1995). The R2 for stepwise
regression models of timing on climate was 0.71 0.09 (mean SD) for plants but only 0.45
0.11 for animals. Second, the predicted advance (in days) in phenology given a realistic future
climate-warming scenario was much greater for plants than for animals (Fig. 3c and 3d, data
from table 6 in Sparks and Carey, 1995). Climate warming is predicted to advance plant
phenology by an average of 19.5 0.9 days in response, but that for animals by only 6.6 1.3
days (P 0.001).

These data are derived from a limited set of species at a single site but nonetheless
support Bradshaw's (1972) general expectation that plants are more tightly coupled to
environmental variation than are animals. The observed pattern might alternatively result from
differential climatic change in the wintering grounds of migratory animals or from the error
associated with defining the return of migrating species (Sparks and Carey, 1995). However,
phenologies of non-migratory animals also showed poor fits with climate data as well as showed
limited predicted response to climate change.
These phenological patterns raise interesting suggestions regarding potential changes in the
distribution and abundance of plant and animal populations in response to climate change. For
example, climate warming might have a bigger impact on the intertidal zonation of sessile than
of mobile species. Whether climate warming is having a bigger impact on plants than on animals
in terrestrial systems remains to be determined; however, distributions of plants and animals
shifted dramatically and individualistically following de-glaciation in the late Quaternary
(Graham and Grimm, 1990).

Do plants and animals differ in macro-evolutionary responses to the environment (Niklas, 1997)?
If plants and animals are indeed sensitive to different kinds of stresses or at least have different
sensitivities, then the timing of their mass extinctions might differ.

The timing of mass extinctions has been studied intensively. Five mass extinctions have
generally been recognized for marine invertebrates (Jablonski, 1991), and two mass extinctions
(plus some minor ones) are recognized for terrestrial vertebrates (Benton, 1989). However, only
three mass extinctions (late Ordovician, Permian-Triassic, Cretaceous-Tertiary) are currently
receiving strong support (R. Bambach and A. Knoll, reported in Kerr, 2001). What about
plants? Niklas (1997) states that nine episodes of intense species extinction characterize the
history of vascular land plants and that none of these extinctions coincides with any of the major
extinctions (post-Silurian) of marine invertebrates or of terrestrial vertebrates (see fig. 8.14
in Niklas, 1997). However, recent data suggests that plants actually experienced only two major
extinctions (Permian-Triasic, Cretaceous-Tertiary; A. H. Knoll, personal communication), which
are in fact synchronous with the mass extinctions of marine and terrestrial animals. Given that
several mass extinctions have recently gone extinct (e.g.,Kerr, 2001), it now seems premature
to evaluate whether extinction patterns of plants and animals actually show differential
sensitivity to environmental changes.

CONCLUDING REMARKS

We have examined some physiological, ecological, and evolutionary consequences of one

fundamental difference between animals and plants, namely, the fact that animals (but not plants)
can use their behavior and mobility to evade environmental stresses and to maximize time in
suitable environments. Our review is based largely on a classic synthesis by A. D. Bradshaw
(1972), in which he drew attention to the evolutionary consequences of various key biological
differences (not just behavioral ones) between plants and animals.

The generalizations given here are largely well known, but nevertheless are worth repeating.
Most importantly, behavior and mobility can be an effective defense against diverse biotic and
environmental stresses. Consequently, any organismbe it a plant or a sessile invertebratethat
has limited capacities in this regard may need to amplify evolutionarily its use of other
mechanisms (e.g., greater tolerance, plasticity, or repair capacity) to deal with stresses
(Bradshaw, 1972). However, comparing quantitative responses of plants and animals has proven
remarkably difficult.

We would be remiss not to reiterate that plants and animals differ, of course, in many ways other
than in behavioral and locomotor capacities (Bradshaw, 1972). For example, they differ in
dispersal distance (Bradshaw, 1972), breeding system (Lande and Schemske, 1985), modular
construction (Harper, 1977, 1986), and ploidy (Ramsey and Schemske, 1998), to say nothing of
their vast physiological differences. All of these differences should have major ecological and
evolutionary consequences (Bradshaw, 1972; Harper, 1977, 1986), will undoubtedly interact
with differences in behavior and locomotion, and should be well worthy of detailed
investigation. Someone interested in embarking on such an investigation would be well advised
to start with a careful reading of Bradshaw (1972).

TABLE 1. Number of citations (Biosis, 1991 to fall 2000) that use combinations of
key words (phenotypic plasticity + plant or animals + one of several environmental
variables).*

View Large

TABLE 2. Results of ANCOVA on the relationship between upper distributional

limits, maximum mid-intertidal temperature, and functional group (algae, sessile
invertebrate, or mobile invertebrate).*
 View Large

View largeDownload slide

FIG. 1. Strength of directional selection on morphological traits of various taxa. (a) The proportion of estimates of
non-mortality selection on plants (solid line, n = 344 estimates), vertebrates (dashed line, n = 234), and insects
(dotted line, n = 308) as a function of the absolute standardized selection gradient (||). (b) The proportion of
estimates of mortality selection on plants (n = 33), vertebrates (n = 189), and invertebrates (n = 56) as a function of
||

View largeDownload slide

FIG. 2. Regression lines for the relationship between mid-intertidal rock temperatures and the upper limits of
representative algae, sessile invertebrates, and mobile invertebrates. Both algae and sessile inverts display sharp
decreases in upper limits with increasing thermal stress, whereas mobile invertebrates do not. Rock temperatures
were measured at a standard 1.5 m above mean lower low water (MLLW) with ibutton temperature loggers (Dallas
Semiconductor). Species' upper limits were determined using surveyors equipment. Relative intertidal height is
scaled from zero (MLLW) to one (mean higher high water, MHHW) based on predicted tidal curves. The species
shown are the algae Fucus gardneri (a), Halosaccion glandiforme (b), and Hedophyllum sessile (c),
barnacles Balanus glandula (d) and Semibalanus cariosus (e), mussel Mytilus californianus (f), chiton Katharina
tunicata(g), and sea star Pisaster ochraceus (h)

View largeDownload slide

FIG. 3. Relative phenological sensitivity of (a) plants and (b) animals to climate change over two centuries. Plotted
in are histograms of R2 values (%) for regressions of time of onset of phenological variables on climate variables for
plants (a) and animals (b). Predicted advance (in days) of phenological variables in response to a simulated climate
warming for (c) plants and (d) animals. Data were collected (see Sparks and Carey, 1995) at the Marsham family
estates (England)
1

From the symposium Plant/Animal Physiology presented at the Annual Meeting of the Society
for Integrative and Comparative Biology, 37 January 2001, at Chicago, Illinois.
2

E-mail: hueyrb@u.washington.edu
3

Present address of Margen Carlson: c/o Graduate Support Services Office (Lab I, Rm 3019), The
Evergreen State College, Olympia, Washington 98505
4

Present address of Christopher Harley: Hopkins Marine Station, Oceanview Blvd., Pacific
Grove, California 93950
5

Present address of Anhthu Hoang: Department of Integrative Biology, University of California,

Berkeley, California 94720

We thank M. E. Feder for the invitation to participate in this symposium and for suggesting the
topic. We thank NSF for covering travel expenses (IBN-0097876) and for supporting Huey's
research (DEB9981598 and INT09981780). C. Harley was supported by a NSF predoctoral
fellowship and by a Padilla Bay National Estuarine Research Reserve research assistantship.
Several botanical colleagues have answered our questions and attempted to reeducate us (J.
Ehleringer, S. Kalisz, J. Maron, D. Schemske, S. Tonsor, L. Van Volkenburgh, H. Young), but
should not be held responsibility for our learning disabilities. J. Ruesink provided analytical
advice. J. Henderson, A. Knoll, R. Lee, and M. Benton answered various questions. We thank
Dan Lewis, who helped us hunt down the delightful letter from C. Darwin to E. Krause: this item
is reproduced by permission of The Huntington Library, San Marino, California. Finally, we
thank D. Schemske for steering us to Bradshaw's paper, and of course A. D. Bradshaw for
writing it.

Land animals

Reindeer are expected to disappear from large portions of their current range by the
end of the century.

Marmots are ending their hibernations about three weeks earlier than they did 30 years
ago.

Canadian red squirrels are breeding about 18 days earlier.

Red foxes are spreading northward, encroaching on territory normally occupied by their
artic cousins.
North American Fowler's toads are breeding six days later than they did a decade ago.

Polar bears today are thinner and less healthy than those of 20 years ago.

Sealife

Coral reefs around the world are predicted to increase by up to a third in size.

Elephant seal pups are leaner because their prey is migrating to cooler waters.

Loggerhead sea turtles are laying their eggs about 10 days earlier than they did 15
years ago.

Rising temperatures are influencing the sex of Hawkbill turtle hatchlings, with more
females than males being born.

Tidal organisms like rock barnacles, mollusks, and tidal snails commonly found in warm
southern waters are moving northward.

Many fish species are moving northward in search of cooler waters.

Birds

The diet of some songbirds are changing, with some avoiding insects that consume
leaves exposed to high levels of carbon dioxide.

North American tree swallows are laying their eggs about nine days earlier than they did
40 years ago.

Common murres are breeding 24 days earlier than they did a decade ago.

Plants

Some plants are thriving in areas where their growth was limited before, thanks to
temperature changes that provide more water, heat and sunlight.

American flowering plants like columbines and wild geraniums are blooming earlier than
before.

Insects

Edith's checkerspot butterflies are moving northward in search of cooler temperatures.

A gene in the fruitfly Drosophila normally associated with hot, dry conditions has spread
to populations living in traditionally cooler southern regions.
Impacts of climate change on individual species are increasingly well documented, but we lack
understanding of how these effects propagate through ecological communities. Here we combine
species distribution models with ecological network analyses to test potential impacts of climate
change on >700 plant and animal species in pollination and seed-dispersal networks from central
Europe. We discover that animal species that interact with a low diversity of plant species have
narrow climatic niches and are most vulnerable to climate change. In contrast, biotic
specialization of plants is not related to climatic niche breadth and vulnerability. A simulation
model incorporating different scenarios of species coextinction and capacities for partner
switches shows that projected plant extinctions under climate change are more likely to trigger
animal coextinctions than vice versa. This result demonstrates that impacts of climate change on
biodiversity can be amplified via extinction cascades from plants to animals in ecological
networks.

Climate change forces species either to move or to adapt to changing conditions1,2. Although
models predicting the responses of individual species to climate change are widely utilized2, it is
not yet clear to what extent a changing climate will affect biotic interactions between species3,4.
Ecological theory predicts that abundant generalist species tend to have large ranges5 and,
consequently, occupy wide climatic niches6, whereas species specialized on specific interaction
partners have small ranges, occupy narrow climatic niches and may therefore be particularly
vulnerable to climate change7.

In ecological communities, species are embedded in networks of interacting species, for instance,
in mutualistic networks between plant species and animal pollinators or seed dispersers8. Species
in these networks vary in the number of interaction partners, for example, because of differences
in species traits9, and thus differ in their degree of biotic specialization8. So far, it has not been
tested how biotic specialization in ecological networks relates to a species climatic niche
breadth and its vulnerability to climate change. However, a quantitative understanding of this
relationship is required to predict the likelihood of species extinctions and coextinctions from
ecological communities under climate change10.

Here we test the two hypotheses that plants and animals with (1) narrow climatic niches and (2) a
projected loss in climatic suitability are biotic specialists that interact with a low diversity of
partners. We additionally simulate (3) how the relationship between biotic specialization and
vulnerability to climate change affects the risk of species coextinctions of plants and animals
under future climatic conditions. We analysed data on climatic niche breadth for 295 species of
plants and their insect pollinators (196 bee, 70 butterfly and 97 hoverfly species) and seed
dispersers (51 bird species) from central Europe. For each species, we quantified the change in
climatic suitability across a species current European range under projected climate change
according to two circulation models and two representative concentration pathways (RCPs 6.0
and 8.5). We linked projected changes in climatic suitability to data on biotic specialization
derived from 8 quantitative pollination and 5 quantitative seed-dispersal networks recorded in 13
regions across central Europe. Networks describe interaction frequencies between plant and
animal species, that is, the number of visits of an animal to a plant species, and yield empirical
estimates of biotic specialization for each species in each network.

We find that animal species with narrow climatic niches and a projected loss in climatic
suitability interact with a low diversity of plant partners, whereas we do not find analogous
relationships for plants. This important difference between plant and animal species affects the
likelihood of species coextinctions under climate change. We simulate different scenarios of
species coextinction and capacities for partner switches and show with these simulations that
mutualistic networks are more sensitive to projected plant than to animal extinctions under
climate change. We conclude that a high potential for adaptive partner switches is required to
stabilize mutualistic networks against extinction cascades from plants to animals under climate
change.

Bacteria and Archaea and the Cycles of Elements in the Environment (page 1)

(This chapter has 4 pages)

Kenneth Todar, PhD

The most significant effect that the procaryotes, bacteria and archaea, have on their environment is their
underlying ability to recycle the essential elements that make up cells. The earth is a closed system with
limited amounts of certain elements in forms that are utilized by cells. These element are generally acted
upon first by microbes to assimilate them into living matter. The total biomass of microbial cells in the
biosphere, their metabolic diversity, and their persistence in all habitats that support life, guarantee that
microbes will play crucial roles in the transformations and recycling of these elements among all forms of
life.

The table below lists the major elements that make up a typical procaryotic cell (in this case, E. coli). As
expected, over 90 percent of the elemental analysis consists of carbon, hydrogen, oxygen, nitrogen,
phosphorus and sulfur. These are the elements that become combined to form all the biochemicals and
macromolecules that comprise living systems. C, H, O, N, P and S are the constituents of organic material
(An organic compound is a chemical that contains a carbon to hydrogen bond. Organic compounds on earth
are evidence of life. Organic compounds may be symbolized as CH2O, which is the empirical formula for a
sugar such as glucose.) H and O are the constituents of water (H2O), that makes up over 95 percent of the
cell composition. Calcium (Ca++), iron (Fe++), magnesium (Mg++) and potassium (K+) are present as
inorganic salts in the cytoplasm of cells.

Table 1. Major elements, their sources and functions in cells.

% of
Element dry Source Function
weight
organic
Main constituent of cellular
Carbon 50 compounds or
material
CO2
Constituent of cell material
H2O, organic
and cell water; O2 is
Oxygen 20 compounds,
electron acceptor in aerobic
CO2, and O2
respiration
 NH3, NO3, Constituent of amino acids,
Nitrogen 14 organic nucleic acids nucleotides,
compounds, N2 and coenzymes
H2O, organic Main constituent of organic
Hydrogen 8
compounds, H2 compounds and cell water
inorganic Constituent of nucleic acids,
Phosphorus 3 phosphates nucleotides, phospholipids,
(PO4) LPS, teichoic acids
SO4, H2S, S, Constituent of cysteine,
Sulfur 1 organic sulfur methionine, glutathione,
compounds several coenzymes
Main cellular inorganic
Potassium 1 Potassium salts cation and cofactor for
certain enzymes
Inorganic cellular cation,
Magnesium
Magnesium 0.5 cofactor for certain
salts
enzymatic reactions
Inorganic cellular cation,
cofactor for certain
Calcium 0.5 Calcium salts
enzymes and a component
of endospores
Component of cytochromes
and certain nonheme iron-
Iron 0.2 Iron salts
proteins and a cofactor for
some enzymatic reactions

The table ignores the occurrence of "trace elements" in cells. Trace elements are metal ions required in
cellular nutrition in such small amounts that it is difficult to determine or demonstrate their presence in cells.
The usual metals that qualify as trace elements are Mn++, Co++, Zn++, Cu++ and Mo++. Trace elements
are usually built into vitamins and enzymes. For example, vitamin B12 contains cobalt (Co++) and the
bacterial nitrogenase enzyme contains molybdenum (Mo++).

Microbes and the Cycles of Elements

Of course, all living organisms play a role in the cycles of elements, but for the most part, it is the
procaryotes that play major and sometimes unique roles. Herein, we discuss total microbial contribution to
the cycles of the major elements, but major emphasis is placed on procaryotes.

The fungi (molds and yeasts). The molds are aerobic organisms that utilize organic compounds for
growth. They play an important role in decomposition or biodegradation of organic matter, particularly in
soil. Yeast can grow anaerobically (without oxygen) through the process of fermentation. They play a role in
fermentations in environments high in sugar. The prominent role of fungi in the environment is in the carbon
cycle, during the process of decomposition, especially in the soil.

The algae are also an important part of the carbon cycle. They are the predominant photosynthetic
organisms in many aquatic environments. The algae are autotrophs, which means they use carbon dioxide
(CO2) as a source of carbon for growth. Hence they convert atmospheric CO2 into organic material (i.e.,
algal cells). Algae also play a role in the oxygen (O2) cycle since their style of photosynthesis, similar to
plants, produces O2 in the atmosphere. Thecyanobacteria are a group of procaryotic microbes, as
prevalent as algae, that have this type of metabolism. Photosynthetic algae and cyanobacteria can be found
in most environments where there is moisture and light. They are a major component of marine plankton
which the basis of the food chain in the oceans.

Protozoans are heterotrophic organisms that have to catch or trap their own food. Therefore, they have
developed elaborate mechanisms for movement and acquiring organic food which they can digest. Their
food usually turns out to be bacterial cells, so one might argue that they are ecological predators that keep
bacterial populations under control in soil, aquatic environments, intestinal tracts of animals, and many
other environments.

The procaryotic bacteria and archaea, as a result of their diversity and unique types of metabolism, are
involved in the cycles of virtually all essential elements. In two cases, methanogenesis (conversion of
carbon dioxide into methane) and nitrogen fixation (conversion of nitrogen in the atmosphere into biological
nitrogen) are unique to procaryotes and earns them their "essential role" in the carbon and nitrogen cycles.

There are other metabolic processes that are unique, or nearly so, in the procaryotes that bear significantly
on the cycles of elements. For example, procaryotes called lithotrophs use inorganic compounds like
ammonia and hydrogen sulfide as a source of energy, and others called anaerobic respirersuse nitrate
(NO3) or sulfate (SO4) in the place of oxygen, so they can respire without air. Most of the archaea are
lithotrophs that use hydrogen (H2) or hydrogen sulfide (H2S) as a source of energy, while many soil bacteria
are anaerobic respirers that can use their efficient respiratory metabolism in the absence of O2.

The basic processes of heterotrophy are spread throughout the bacteria. Most of the bacteria in the soil and
water, and in associations with animals and plants, are heterotrophs. Heterotrophy means living off of
dead organic matter, usually by some means of respiration (same as animals) or fermentation (same as
yeast or lactic acid bacteria). Bacterial heterotrophs in the carbon chain are important in the processes of
biodegradation and decomposition under aerobic and anaerobic conditions.

In bacteria, there is a unique type of photosynthesis that does not use H2O or produce O2 which impacts on
the carbon and sulfur cycles.

Meanwhile, the cyanobacteria (mentioned above) fix CO2 and produce O2 during photosynthesis, and they
make a very large contribution to the carbon and oxygen cycles.

The list of examples of microbial involvement in the cycles of elements that make up living systems is
endless, and probably every microbe in the web is involved in an intimate and unique way.

The Oxygen Cycle

Basically, O2 is derived from the photolysis of H2O during plant (oxygenic) photosynthesis. Two major
groups of microorganisms are involved in this process, the eucaryotic algae, and the procaryotic
cyanobacteria (formerly known as "blue-green algae"). The cyanobacteria and algae are the source of much
of the O2 in the earth's atmosphere. Of course, plants account for some O2 production as well, but the
microbes predominate in marine habitats which cover the majority of the planet.

Since most aerobic organisms need the O2 that results from plant photosynthesis, this establishes
a relationship between plant photosynthesis and aerobic respiration, two prominent types of metabolism on
earth. Photosynthesis produces O2 needed for aerobic respiration. Respiration produces CO2 needed for
autotrophic growth.

CO2 + H2O-----------------> CH2O (organic material) + O2 plant (oxygenic) photosynthesis

CH2O + O2-----------------> CO2 + H2O aerobic respiration

Since these photosynthetic microbes are also autotrophic (meaning they convert CO2 to organic material
during growth) they have a similar impact on the carbon cycle (page 2).
Bacteria and Archaea and the Cycles of Elements in the Environment (page 2)

(This chapter has 4 pages)

Kenneth Todar, PhD

The Carbon Cycle

Carbon is the backbone of all organic molecules and is the most prevalent element in cellular (organic)
material. In its most oxidized form, CO2, it can be viewed as an "inorganic" molecule (no C - H
bond). Autotrophs, which include plants, algae, photosynthetic bacteria, lithotrophs, and methanogens, use
CO2 as a sole source of carbon for growth, which reduces the molecule to organic cell material
(CH2O). Heterotrophs require organic carbon for growth, and ultimately convert it back to CO2.

Thus, a relationship between autotrophs and heterotrophs is established wherein autotrophs fix carbon
needed by heterotrophs, and heterotrophs produce CO2 used by the autotrophs.

CO2 + H2O-----------------> CH2O (organic material) autotrophy

CH2O + O2-----------------> CO2 + H2O heterotrophy

Since CO2 is the most prevalent greenhouse gas in the atmosphere, it isn't good if these two equations to
get out of balance (i.e. heterotrophy predominating over autotrophy, as when rain forests are destroyed and
replaced with cattle).

Autotrophs are referred to as primary producers at the "bottom of the food chain" because they convert
carbon to a form required by heterotrophs. Among procaryotes, the cyanobacteria, the lithotrophs and the
methanogens are a formidable biomass of autotrophs that account for a corresponding amount of
CO2fixation in the global carbon cycle.

The lithotrophic bacteria and archaea that oxidize reduced N and S compounds and play important roles in
the natural cycles of N and S (discussed below), are virtually all autotrophs. The prevalence of these
organisms in sulfur-rich environments (marine sediments, thermal vents, hot springs, endosymbionts, etc.
may indicate an unappreciated role of these procaryotes as primary producers of organic carbon on earth.

The methanogens play a dual role in the carbon cycle. These archaea are inhabitants of virtually all
anaerobic environments in nature where CO2 and H2(hydrogen gas) occur. They use CO2 in their metabolism
in two distinct ways. About 5 percent of CO2 taken up is reduced to cell material during autotrophic growth;
the remaining 95 percent is reduced to CH4 (methane gas) during a unique process of generating cellular
energy. Hence, methane accumulates in rocks as fossil fuel ("natural gas"), in the rumen of cows and guts
of termites, in sediments, swamps, landfills and sewage digesters. Since CH4 is the second-most prevalent
of the greenhouse gases, it is best to discourage processes that lead to its accumulation in the atmosphere.

CO2 + H2 -----------------> CH2O (cell material) + CH4 methanogenesis

Under aerobic conditions, methane and its derivatives (methanol, formaldehyde, etc.) can be oxidized as
energy sources by bacteria called methylotrophs.Metabolically this is a version of decomposition or
biodegradation during the carbon cycle which is discussed below.

Biodegradation is the process in the carbon cycle for which microbes get most credit (or
blame). Biodegradation is the decomposition of organic material (CH2O) back to CO2 + H2O and H2. In
soil habitats, the fungi play a significant role in biodegradation, but the procaryotes are equally important.
The typical decomposition scenario involves the initial degradation of biopolymers (cellulose, lignin, proteins,
polysaccharides) by extracellular enzymes, followed by oxidation (fermentation or respiration) of the
monomeric subunits. The ultimate end products are CO2, H2O and H2, perhaps some NH3 (ammonia) and
sulfide (H2S), depending on how one views the overall process. These products are scarfed up by lithotrophs
and autotrophs for recycling. Procaryotes which play an important role in biodegradation in nature include
the actinomycetes, clostridia, bacilli, arthrobacters and pseudomonads.

Overall Process of Biodegradation (Decomposition)

polymers (e.g. cellulose)-----------------> monomers (e.g. glucose)depolymerization

monomers-----------------> fatty acids (e.g. lactic acid, acetic acid, propionic acid) + CO2 + H2 fermentation

monomers + O2 -----------------> CO2 + H2O aerobic respiration

The importance of microbes in biodegradation is embodied in the adage that "there is no known natural
compound that cannot be degraded by some microorganism." The proof of the adage is that we aren't up to
our ears in whatever it is that couldn't be degraded in the last 3.5 billion years. Actually, we are up to our
ears in cellulose and lignin, which is better than concrete, and some places are getting up to their ears in
teflon, plastic, styrofoam, insecticides, pesticides and poisons that are degraded slowly by microbes, or not
at all.

Figure 1. The Carbon Cycle. Organic matter (CH2O) derived from photosynthesis (plants, algae and
cyanobacteria) provides nutrition for heterotrophs (e.g. animals and associated bacteria), which convert it
back to CO2. Organic wastes, as well as dead organic matter in the soil and water, are ultimately broken down
to CO2 by microbial processes of biodegradation.

The figure above mostly ignores the role of methanogenesis in the carbon cycle. Since methanogens have
the potential to remove CO2 from the atmosphere, converting it to cell material and CH4, these procaryotes
not only influence the carbon cycle, but their metabolism also affects the concentration of major greenhouse
gases in earth's atmosphere.

Recently, I asked a colleague, Professor Paul Weimer of the University of Wisconsin Department of
Bacteriology, whether mathanogenesis, which utilizes CO2 while producing CH4 was better or worse on the
greenhouse effect and global warming. This is his response. "Worse. During methanogenesis involving
CO2 reduction, the stoichiometry is 4H2 + CO2 --> CH4 + 2 H2O, so one mole of a greenhouse gas is
exchanged for another. But methane is about 15 times more potent than is CO2 in terms of heat absorption
capability on a per-molecule basis, so the net effect is a functional increase in heat absorption by the
atmosphere. Remember also that in most natural environments, around two-thirds of the methane is
produced by aceticlastic methanogenesis (CH3COOH --> CH4 + CO2) - an even less welcome situation, as
BOTH products are greenhouse gases.

Even though methane concentrations in the atmosphere are two orders of magnitude below those of CO2,
methane is thought to account for about 15% of the anthropogenic climate forcing, compared to about 60%
from CO2. Most of the rest of the contribution is from nitrous oxide (N2O, a respiratory denitrification
product that has something like 300 times the heat absorbing capacity as CO2) and the old
chlorofluorocarbons (CFCs), even stronger heat absorbers yet, but more famous and dangerous as
stratospheric ozone-depleters."
Bacteria and Archaea and the Cycles of Elements in the Environment (page 3)

(This chapter has 4 pages)

Kenneth Todar, PhD

The Nitrogen Cycle

The nitrogen cycle is the most complex of the cycles of elements that make up biological systems. This is
due to the importance and prevalence of N in cellular metabolism, the diversity of types of nitrogen
metabolism, and the existence of the element in so many forms. Procaryotes are essentially involved in the
biological nitrogen cycle in three unique processes.

Nitrogen Fixation: this process converts N2 in the atmosphere into NH3(ammonia), which is assimilated
into amino acids and proteins. Nitrogen fixation occurs in many free-living bacteria such as clostridia,
azotobacters and cyanobacteria, and in symbiotic bacteria such as Rhizobium and Frankia, which associate
with plant roots to form characteristic nodules. Biological nitrogen fixation is the most important way that
N2 from the air enters into biological systems.

N2 ----------------> 2 NH3 nitrogen fixation

Anaerobic Respiration: this relates to the use of oxidized forms of nitrogen (NO3and NO2) as final electron
acceptors for respiration. Anaerobic respirers such as Bacillus and Pseudomonads are common soil
inhabitants that will use nitrate (NO3) as an electron acceptor. NO3 is reduced to NO2 (nitrite) and then to a
gaseous form of nitrogen such as N2 or N2O (nitrous oxide). The process is called denitrification. (A related
process conducted by some Bacillus species, called dissimilatory nitrate reduction reduces NO3 to
ammonia (NH3), but this is not considered denitrification.) Denitrifying bacteria are typically facultative
microbes that respire whenever oxygen is available by aerobic respiration. If O2 is unavailable for
respiration, they will turn to the alternative anaerobic respiration which uses NO3. Since NO3 is a common
and expensive form of fertilizer in soils, denitrification may not be so good for agriculture, and one rationale
for tilling the soil is to keep it aerobic, thereby preserving nitrate fertilizer in the soil.

NO3 ----------------> NO2 ----------------> N2 denitrification

The overall reactions of denitrification shown above proceed through the formation of nitrous oxide (N2O). A
recent article by Wunsch an Zumft in Journal of Bacteriology, vol. 187 (2005), sheds new light on the process
of denitrification. N2O is a bacterial metabolite in the REVERSAL of Nitrogen fixation. The anthropogenic
atmospheric increase of N2O is a cause for concern, as noted above (as a greenhouse gas, N2O has 300 times
the heat absorbing capacity as CO2). Denitrifying bacteria respire using N2O as an electron acceptor yielding
N2 and the thereby provide a sink for N2O. This article provides new insight into this process by identifying a
membrane-bound protein in denitrifying bacteria called NosR, that is necessary for the expression of N 2O
reductase from the nosZ gene. The NosR protein has redox centers positioned on opposite sides of the
cytoplasmic membrane, which allows it to sustain whole-cell N2O respiration by acting on N2O reductase.
Nitrification is a form of lithotrophic metabolism that is chemically the opposite of denitrification. Nitrifying
bacteria such as Nitrosomonas utilize NH3 as an energy source, oxidizing it to NO2, while Nitrobacter will
oxidize NO2 to NO3. Nitrifying bacteria generally occur in aquatic environments and their significance in soil
fertility and the global nitrogen cycle is not well understood.

The Overall process of Nitrification

NH3 ----------------> NO2 (Nitrosomonas)

NO2 ----------------> NO3 (Nitrobacter)

A final important aspect of the nitrogen cycle that involves procaryotes, though not exclusively,
is decomposition of nitrogen-containing compounds. Most organic nitrogen (in protein, for example) yields
ammonia (NH3) during the process of deamination. Fungi are involved in decomposition, as well.

Plants, animals and protista, as well as the procaryotes, complete the nitrogen cycle during the uptake of
the element for their own nutrition. Nitrogen assimilation is usually in the form of nitrate, an amino group,
or ammonia.

Figure 2. The Nitrogen Cycle

Bacteria and Archaea and the Cycles of Elements in the Environment (page 3)

(This chapter has 4 pages)

Kenneth Todar, PhD

The Nitrogen Cycle

The nitrogen cycle is the most complex of the cycles of elements that make up biological systems. This is
due to the importance and prevalence of N in cellular metabolism, the diversity of types of nitrogen
metabolism, and the existence of the element in so many forms. Procaryotes are essentially involved in the
biological nitrogen cycle in three unique processes.

Nitrogen Fixation: this process converts N2 in the atmosphere into NH3(ammonia), which is assimilated
into amino acids and proteins. Nitrogen fixation occurs in many free-living bacteria such as clostridia,
azotobacters and cyanobacteria, and in symbiotic bacteria such as Rhizobium and Frankia, which associate
with plant roots to form characteristic nodules. Biological nitrogen fixation is the most important way that
N2 from the air enters into biological systems.

N2 ----------------> 2 NH3 nitrogen fixation

Anaerobic Respiration: this relates to the use of oxidized forms of nitrogen (NO3and NO2) as final electron
acceptors for respiration. Anaerobic respirers such as Bacillus and Pseudomonads are common soil
inhabitants that will use nitrate (NO3) as an electron acceptor. NO3 is reduced to NO2 (nitrite) and then to a
gaseous form of nitrogen such as N2 or N2O (nitrous oxide). The process is called denitrification. (A related
process conducted by some Bacillus species, called dissimilatory nitrate reduction reduces NO3 to
ammonia (NH3), but this is not considered denitrification.) Denitrifying bacteria are typically facultative
microbes that respire whenever oxygen is available by aerobic respiration. If O2 is unavailable for
respiration, they will turn to the alternative anaerobic respiration which uses NO3. Since NO3 is a common
and expensive form of fertilizer in soils, denitrification may not be so good for agriculture, and one rationale
for tilling the soil is to keep it aerobic, thereby preserving nitrate fertilizer in the soil.

NO3 ----------------> NO2 ----------------> N2 denitrification

The overall reactions of denitrification shown above proceed through the formation of nitrous oxide (N2O). A
recent article by Wunsch an Zumft in Journal of Bacteriology, vol. 187 (2005), sheds new light on the process
of denitrification. N2O is a bacterial metabolite in the REVERSAL of Nitrogen fixation. The anthropogenic
atmospheric increase of N2O is a cause for concern, as noted above (as a greenhouse gas, N2O has 300 times
the heat absorbing capacity as CO2). Denitrifying bacteria respire using N2O as an electron acceptor yielding
N2 and the thereby provide a sink for N2O. This article provides new insight into this process by identifying a
membrane-bound protein in denitrifying bacteria called NosR, that is necessary for the expression of N 2O
reductase from the nosZ gene. The NosR protein has redox centers positioned on opposite sides of the
cytoplasmic membrane, which allows it to sustain whole-cell N2O respiration by acting on N2O reductase.

Nitrification is a form of lithotrophic metabolism that is chemically the opposite of denitrification. Nitrifying
bacteria such as Nitrosomonas utilize NH3 as an energy source, oxidizing it to NO2, while Nitrobacter will
oxidize NO2 to NO3. Nitrifying bacteria generally occur in aquatic environments and their significance in soil
fertility and the global nitrogen cycle is not well understood.

The Overall process of Nitrification

NH3 ----------------> NO2 (Nitrosomonas)

NO2 ----------------> NO3 (Nitrobacter)

A final important aspect of the nitrogen cycle that involves procaryotes, though not exclusively,
is decomposition of nitrogen-containing compounds. Most organic nitrogen (in protein, for example) yields
ammonia (NH3) during the process of deamination. Fungi are involved in decomposition, as well.

Plants, animals and protista, as well as the procaryotes, complete the nitrogen cycle during the uptake of
the element for their own nutrition. Nitrogen assimilation is usually in the form of nitrate, an amino group,
or ammonia.
Figure 2. The Nitrogen Cycle