Tolerance Range

We refer to the surroundings of an organism as its environment or habitat. The basic subdivision is
between terrestrial, or land, habitats and aquatic, or watery, habitats. A trout in aquatic organism,
living in cool streams, whereas a trout lily is terrestrial, living in moist forests. Aquatic habitats are
often subdivided into marine, or ocean, habitats, and fresh-water habitats which are lakes, ponds,
streams, and springs whose water usually have low salt concentrations. Much finer subdivisions can
be made-habitats may be subdivided into sub habitats and eventually microhabitats, such as a
south-facing slope, a tree hole, or the spaces between the sand grains along a beach.
Specific features of the habitat may be studied. For the trout we can measure such factors as water
temperature and the amount of oxygen dissolved in the water. For the trout lily we can measure the
amount of light reaching it on the forest floor. And the concentration of calcium in the soil. If we are
able to determine the whole range over which the species is able to live, for one of these factors, we
then will know the range of tolerance of that species for that factor. A goldfish, for example, might
be able to live in water in which the temperature ranges from 2 to 34oC (about 36 to 39oF), but it will
die of heat or cold at temperature above or below this range.
Field observations give us some idea of the range of tolerance of a species, but the field work must
be checked by experimentation in the laboratory and in the field. One of the reasons for this is that
organisms interact with other organisms. The habitat feature we have mentioned so far have all
been abiotic factors, or factor of the physical environment. The organism also exists in world of
biotic factors, consisting of its relationship with other organisms. These relationships are of several
kinds: one organism may use another as food and at the same time serve as food for the third; two
organisms may compete for food, for nesting sites, or for some other requirement. An organism may
not be able to live in an area where physical factors are favorable for it due to some biotic factors.
Plants that tent to be found growing only in acid soils are called calciphobes. Some of these
specimen may do so because they are unable to tolerate neutral or basic soils but this is not always
the case. In this ecology lectures (I have been told), Henry Chandler Cowles described a visit to the
botanical garden in Berlin, which had an excellent collection of calciphobes and their opposite
number, calciphiles, characteristic of basic soils. Cowls questioned one of the gardeners, asking if it
wasnt difficult treating the soil in the different beds so as to maintain each at just the right pH. Not
at all, the gardener is supposed to have answered; we grow them all in the same soil.
Clearly such species in nature are being restricted to just one end of their tolerance ranges.
Competition from the many species that do well in average favorable condition is probably often the
cause of such restriction to extreme habitats.
This is not say that pH or other abiotic factors do not limit plant or animal distribution. Often, the
apparent physical cause of an organisms restriction is the real one. The absence of lichens century,
and research since then has tended to confirm that sensitivity to sulfur dioxide is the main factor in
this distribution (Anderson and Treshow 1984).
In other case, the apparent abiotic causes are not borne out when experiments are done. An annual
plant called sea rocket along the Pacific Coast of the United States grows only on open, bare sandy
beaches in the salt-spray zone. When M. G. Barbour and his coworkers (1973) at the University of
California at Davis planted seeds in various habitats, they found that the seedlings grew best inland
in good grassland soil (Figure 2-1). This was true, however, only when the area of grassland soil was
weeded of competitors; in undisturbed grassland, another biotic factor came into play; grazing by
small mammals was severe on the sea rockets growing on the grassland site.

Tolerance range differ from one species to another. We would expect carp (goldfish), for example, to
be able to tolerate warmer temperatures and lower oxygen concentrations than trout, and this is
exactly what we find when we make appropriate test (Figure 2-2). The fact that different species
have different tolerance ranges is one basic reason why different habitats support different
communities, and why communities vary geographically from warmer to cooler and from wetter to
drier regions
Range of the optimum
Generally, an organism does not do equally well throughout its whole range of tolerance. Instead
there is some part, usually toward the middle, where it does better (Figure 2-3). This section is
referred to as the range of optimum. If we look at the rate of any one process at different levels of
any one factors, we can usually specify optimum range with no trouble. For example, we may find
that pea plants photosynthesize fastest around 30oC. Does this mean that this is the optimal
temperature? It is for photosynthesize, but it may not be the most favorable temperature for
another function, such us growth. Photosynthetic rate and respiration rate both go up with
temperature, but photosynthetic rate drops at higher temperatures whereas respiration rate keeps
increasing. When this two curves are combined, the temperature at which growth is greatest will be
below 30oC, perhaps 25oC (Figure 2-4).
So, is 25oC the optimal temperature for pea plants? They grow fastest at that temperature, but is
this the temperature at which they reproduce fastest or live longest? Possibly not. Quite different
temperatures may be optimal for growth and for flowering, for example. Even considering just
growth rate, the example is oversimplified because the effects of day-night temperature changed
(thermoperiodism) are not considered.
The ecological optimum is a useful concept but a blunt one, like a hammer. When precision is
needed, some other tool may be better.
When condition change
Few habitats stay the same for very long. Some, such as the ocean depths, come close, but for most
habitats environmental factors change between day and night, between drought and wet period,
between summer and winter. When some important feature of the habitat changes, the organism
changes in response. The change may be mainly physiological. As the air temperature warms up
during the day, a plant and a bird both exposed to the sun will undergo physiological changes
because of the heat. These changes often have effect of keeping certain important aspects of the
organisms internal environment. This tendency, which is an important physiological principle, is
known as homeostatis. The bird may respond by lowering its rate of heat production and by
arranging its feathers so as to long heat rapidly, resulting in its body temperature remaining
constant.
If the sun keeps shining and the temperature keeps rising, an important difference between the
plant and the bird or the trout lily and the trout may be seen. The plants are stationary, attached to
one spot (sessile), whereas the animals can move about (motile). The bird or trout can move away
from unfavorable conditions-the bird can go into the shade; the trout can swim to cooler water.
These are behavioral changes that occur in addition to the purely physiological ones. The range of
behavioral responses for a sessile organism is much smaller.
Phenotypic ways of coping with changed environments

If some environment factor seasonally shifts beyond the tolerance range of an organism, it can do
various things:
1. It can fail to cope, in which case it dies. In this case, the species disappears from the biota of
the area unless it reinvades.
2. The adult can die but some other life history stage, such as eggs, pupae, or seeds, with
broader tolerance limits survives. This strategy can provide for the regular yearly of the
forest and for annual herbs. It may also tide organism over a longer period of unfavorability.
For example, many plants produce seeds that remain viable in the soil for years or even
decades (termed a seed banks) (Leck et al, 1989). An open-country plant may persist in the
seed bank of a site made uninhabitable by the growth of trees and reappear if the trees are
cut or burned.
3. It can hibernate or enter some other type of resting state (Storey and Storey 1990). It can
move to a protected spot below the frost line in the soil where the temperature is low
enough that the animal has a slow, energy-saving metabolism but not low enough to kill it.
Many snakes, turtles, frogs, woodchucks, and invertebrates cope with cold seasons by
hibernation. Aestivation is a similar inactive state shown by various animals in response to
intolerably bot or dry coditions.
4. It can migrate. Many birds live in one geographical area in the summer and another in
winter; so do few mammals and insects. The scale of migration varies enormously. Olivesided flycatchers may migrate from Canada to Ecuador, whereas some soil invertebrates
merely move below the forest line.
5. It can acclimate.
This last way of coping with environmental change is less a part of common knowledge than
hibernation and migration. The range of temperature tolerance of a fish (or an insect or a pine tree)
may vary from season to season, depending of the temperature at which it has recently been living
(Figure 2-5). In midwinter, a fish may be able to live in the range from 0 to 24oC, but down only to
15oC. Such basically physiological adjustments to a changed environment are known as acclimation.
Acclimation does not occur just in temperature tolerance; organism may also acclimate to different
oxygen levels in the air (if they go to higher altitudes, for example) and to other changed conditions.
Acclimation is of great importance in allowing organism to exist permanently in changeable
environments.
Phenotypic Plasticity
In the long term, evolutionary changes may occur in organisms as a consequence of changed
environments. The first installment of our discussion of evolutionary change can be found in a later
part of this chapter and the rest in the population ecology chapters. In the preceding section, we
talked about one of type of non-evolutionary adaptive change in individuals the physiological
changes referred to as acclimation. Acclimation is one category of a broad group of non-evolutionary
changes included under the heading of phenotypic plasticity.
Phenotypic plasticity is defined as environmentally induced phenotypic variation (Stearns 1989).
Phenotypic plasticity, then encompasses all the sorts of differences that can be shown by an
individual (or by several individuals of one genotype, such as a clone) as a result of environment.
In some cases, the resulting variation in discontinuous, such as the production of two different
morphological, physiological, or life history types depending on conditions. A striking example of this
type of phenotypic plasticity involves a moth feeding on oaks (Stearns 1989). Eggs hatching in the

spring give rise to caterpillars that feed on catkins, encounter low levels of tannins, and develop
traits that camouflage them among the catkins. Eggs hatching a little later feed on leaves, encounter
higher levels of tannins, and develop traits that camouflage them on oak leaves. Evidently tannin
level in the food is the developmental switch that leads to the two different morphological types.
One well-studied type of phenotypic plasticity involves the morphological changes of various
planktonic animals, such as water fleas (cladocerans) in response to the presence of predators, or
sometimes just to the changes in physical factors that will usually correlate with the presence of
abundant predators (Dodson 1989). Under these conditions, newly hatched zooplankters grow up to
have various sorts of spines or armaments that discourage capture or ingestion by the predators
(such as copepods or midge fly larvae).
If the change in phenotype with environment is continuous, the relationship is often characterized
using reaction norms. The general idea of reaction norm is shown in Figure 2-6. An individual of
genotype A develops one phenotype when it grows up under environment 1 and a different
phenotype under environment 2. The environmental features we are graphing may be temperature,
crowding, food, or a variety of other things. The phenotypic feature on other axis could be size, shell
thickness, time to metamorphosis, date of egg laying, etc. if we are dealing with a number of
individuals, there is likely to be a range of environments at each locality and accordingly, a range of
phenotypes, as in Figure 2-7.
If the population contains more than one genotype, there will be a bundle of reaction norm lines
rather than one. The reaction norms for the different genotypes may be parallel, or some may cross,
as in figure 2-8. Where this is the case, the relative rankings of genotypes may shift depending on
environment. Suppose, for example, that the phenotype in Figure 2-8 is size. In environment 1,
genotype B produces the bigger organisms, but in environment 1, genotype A produces bigger ones.
Presumably much of the phenotypic plasticity seen in nature is adaptive; that is, the changes result
in better survival of the organisms. This has occasionally been demonstrated and is logical for many
other cases. It is likely that maintaining a broad tolerance range is costly for an organism in energetic
or genetic terms. Phenotypic plasticity in the form of reaction norms may allow a given individual to
develop a narrow tolerance range that is, nevertheless, adaptive for the environment it is occupying
(Gabriel and Lynch 1992).
Extreme conditions
The ability of many organisms to acclimate to temperature, the salt content of water and various
other factors is one complication that must be dealt with in going from the laboratory to the fieldthat is, in explaining the occurrence of organisms in nature on the basis of experimental findings.
Another fact that must be remembered is that conditions need not be perpetually outside the
tolerance limits of a species for the species to be absent. This was pointed out by W. P. Taylor
(1934), one of the early student wildlife management in the western United States. The point is, you
only have to kill a plant or animal once. Six days a week a stream may be well within the tolerance
limits of a certain fish, but if on Saturday night the factory releases a dose of toxic efficient, the fish
will not be found living there. The occasional extreme condition may be more important than the
average condition in the determining whether or not an organism can exist permanently in a given
area.
Daily fluctuations.

It is well to remember that constant conditions, such as are often maintained in the laboratory, are
abnormal to most organism. In most habitats, organisms are exposed to daily alternations of many
featured related to the change from day to night. This is obviously true of light but, depending on
the habitat, it may also be true of such factors as temperature, humidity, pH, and oxygen
concentration.
Organism may respond differently when they experience the alternation of conditions typical of
their natural habitat instead of artificial constant conditions. Growth may be faster, reproductive
rate may be higher (Hoffman 1947), and a higher percentage of seed may germinate when subjected
to alternating warm and cool temperatures than when kept at either of the temperatures without
alternation (Sabo et al. 1979). Such a physiological or ecological response to alternating
temperatures is called thermoperiodism. An example is given in Figure 2-9.
Some habitats, such as grassland, deserts, and shallow ponds, show very large changes between ay
and night. Certain other habitats may be nearly as content as laboratory chambers. Some of the
more constant natural environments are hot springs, caves, the ocean depths, and the inside of
homoiotherms (as a habitat for parasites like blood flukes).