Biology 20C - Fall 1998

ECOLOGY AND EVOLUTIONARY BIOLOGY

Lecture 17 - Ecology of Individuals

 

An ecological individual is a physiological and ecological unit that responds independently, (or potentially independently) of other individuals to its environment. Such individuals may be:

 

Solitary

(Unitary)

units are unattached

(e.g. mammal, bird, tree, snail, anemone, diatom ...)

Colonial

units are close together or attached, but with little interdependence

(e.g. strawberry, aspen, fungi, bacteria ...)

Modular

units remain attached, and are physiologically interdependent

(e.g. coral polyps, portugese man-of-war, tapeworm, Volvox...

 

A genetic individual (genet) consists of all the tissues derived from a single zygote. In solitary organisms, the ecological individual is the same as the genet. In colonial and modular organisms, the genet is divided into genetically identical ramets (a subset of the tissues derived from a zygote). The ramets may be morphologically, physiologically and ecologically similar (monomorphic e.g. banana plants, coral polyps), or they may differ greatly in form and function (polymorphic e.g. reproductive, defensive, feeding polyps in Hydrozoa; leaves, sepals, petals, carpels, stamens in Angiosperms). Colonial and (to a lesser extent) modular forms may be chimeras, containing ramets from two or more genets (e.g. grafted plants, ascidians).

 

Properties of an Individual:

 

States/Events

vs

Processes

 

Birth

 

fertilization

 

Death

 

 

 

Life

birth è death

development

 

Location

fixed

 

 

 

variable

locomotion, dispersal, migration

 

Size (body)

 

growth (physiological; developmental)

 

Age

 

development

 

Sex

 

reproduction

 

Environments of Individuals:

To survive, grow and complete its life cycle, an individual's environment must include necessary:

Conditions - usually physical and chemical factors remaining within certain limits (e.g. temperature, humidity, pH, salinity …)

Resources - physical, chemical or biological materials that are used or consumed by the individual (e.g. substrates, food, shelter, nesting sites…)

 

Necessary conditions and resources may vary greatly during an individual’s life as its physiological and developmental states change. These changes may be permanent (irreversible) or reversible; if reversible, they may be predictable (e.g. seasonal) or erratic responses (to other factors).

 

Individual's Responses to the Environment:

These may be physiological, behavioral, developmental, anatomical, morphological, or ecological. Many physiological responses exist as differences between internal and external environments:

 

Conformation

Internal environment same as external. Individual's ecology controlled largely by instantaneous environmental conditions

Regulation

Internal environment "buffered"; minimizing more extreme external environmental fluctuations

Homeostasis

Internal environment almost constant. Ecology & physiology largely independent of external fluctuations

 

The progression from conforming è regulating è homeostasis leads to increasing physiological and ecological independence of the individual from the stresses of external environmental changes. But regulation and (especially) homeostasis are increasingly "expensive" in terms of the energy, food and other resources needed to maintain the differences between internal and external environments.

 

Principles of Allocation:

As metabolic "costs" rise, organisms increasingly have internal mechanisms to assign priorities to particular functions, and then to allocate energy etc. to the functions. The basic priority is:

 

1. Survival

Minimum requirements to remain alive (in poor condition)

2. Maintenance

Additional requirements to maintain a healthy body

2. Growth

Requirements to increase body size and continue development

4. Reproduction

Requirements for successful reproduction (= complete life cycle)

 

Conformers tend to have little flexibility in how they use energy and resources: allocation is usually sequential and additive - excess energy/resources are used for the next higher function.

In general, conformers use little energy for survival and maintenance (they simply shut down); instead most resources are devoted to rapid growth and reproduction. Conformers tend to be small-bodied, short-lived, rapidly developing organisms.

 

Regulators and (especially) homeostatic organisms tend to be larger-bodied, longer-lived and slower developers. They tend to devote most energy and resources to survival and maintenance, and much smaller proportions to growth and reproduction. However they often accumulate extensive food and energy reserves (as fat, protein or carbohydrate) that allow them to continue "normal" physiological and ecological activity through prolonged periods of variable and/or adverse conditions. They also have considerable flexibility in how energy is allocated to functions: changing priorities can maximize benefits during favorable conditions, and minimize effects of adverse conditions.

 

Tolerance Response Curves:

Physiological tolerances and responses are measured in the laboratory by exposing individuals to an array of conditions along an environmental gradient (usually one factor, e.g. temperature) and measuring a physiological or ecological index of the individual's performance under that condition (e.g. metabolic rate, fat content, growth rate, number of young). Plotting the index of performance (y-axis) against the intensity of the environmental gradient (x-axis) gives a Response Curve. The range of conditions under which the organism can carry out a specified function defines the Zone of Tolerance for that function along that gradient. A typical response curve is bell-shaped, and the four functions (listed above) tend to have increasingly narrower zones of tolerance that are "nested" within one another, and clustered about the conditions (Optimal or Preferred) where performance is greatest:

 

1. Survival

The broadest zone describing all non-lethal parts of the gradient; in the tails of the curve, survival is the only possible function.

2. Maintenance

Maintaining a healthy body is usually possible only in a narrower zone than that in which survival is possible

3. Growth

Often limited to a relatively narrow zone of benign conditions surrounding the optimum

4. Reproduction

Usually the narrowest zone, close to the optimum, and the only conditions where all four functions can be done successfully

 

Dominant mechanisms for responding to the environment and for allocating resources are often correlated with the shapes of the response curves. Conformers tend to have tall, narrow response curves with acute peaks and spanning only a small part of the gradient. Homeostatic organisms tend to have low, broad, flat-topped response curves spanning a large part of the environmental gradient. Regulators lie somewhere on a continuum between conformer and homeostasis.

 

Environmental Patchiness or Heterogeneity ("Grain")

Grain is an index of the scale of environmental variation (in space and/or time), relative to the scale on which the organism detects ("perceives") and responds to environmental variation. Grain is a continuum with endpoints defined by analogy with sandpaper:

 

Coarse-grained

Environmental patches are "large" relative to the organism's "perception"; it spends long periods under the same conditions (homogeneous habitat)

Fine-grained

Environmental patches are "small" relative to the organism's "perception"; it spends little time under any set of conditions (heterogeneous habitat)

 

Dominant response modes are also often correlated with environmental grain. Conformers tend to be limited to relatively coarse-grained situations - they do very well as long as conditions are close to optimal. Conversely, homeostatic organisms are often successful in fine-grained situations, performing well under a broad range of conditions. Regulators are usually intermediate.

 

Ecological Niche

 The concept of "niche" has been very important in the development of many ecological ideas. The concept itself has evolved over the decades, encompassing at least three major ideas. The development of the niche concept is presented below as a historical progression

  A. The Concept of Niche as "Ecological Place"

This original concept arose directly from extensive physiological studies, usually in the laboratory, of individual's response curves and especially of their limits of tolerance. "Place" means the physical-chemical "space" within which the organism can exist. This approach assumes that the abiotic environment is the major control of ecological processes, and mainly deals with the properties of individuals. It is commonly represented as a :

 

1-dimensional tolerance curve

1 environmental condition (a line)

2- or 3- dimensional tolerance space

2 or 3 environmental conditions (rectangle or box)

n-dimensional hypervolume

Many environmental conditions; a mathematical abstraction that is difficult to show graphically

 

B. The Concept of Niche as "Resource Use"

This concept treats resources (materials used or consumed by organisms) in the same way as abiotic conditions. Each resource is arrayed along a quantitative (e.g. seed size, sugar concentration) or qualitative (kind or seeds) resource spectrum (gradient), and some index of biological performance is measured. Resource use can be represented by 1-, 2-, 3- Ö n-dimensional axes, just like those used for "ecological place". Resource use leads to the process of ecological competition. This concept of niche dominated thinking for several decades during which it was accepted that competition for resources was the major ecological interaction among organisms. This concept is mainly concerned with properties of populations.

 

C. The Concept of Niche as "Ecological Role"

This is the broadest and most recent niche concept, incorporating any kind of ecological interaction with other organisms. "Role" refers to the target organism's part in each interaction (as competitor; predator or prey; parasite, pathogen or host; symbiont or host). Each individual may play many different roles during its life, and different individuals may play different roles. Some roles may continue for long periods, while others may be limited to rare, brief interludes. Role is the most abstract concept of niche, often difficult to quantify, and usually unsuited to graphical representation. It is most relevant when dealing with properties of communities.

 

Modern Concepts and Usage of "Niche"

It is currently fashionable to avoid using the term "niche", but the concepts and thinking associated with this term permeates much modern ecological thinking. Niche may include any or all of the three concepts (space, resources, role), so it is the responsibility of the user to define clearly how "niche" is being used. It may be measured by aspects of any or all of the four functions (survival, maintenance, growth, reproduction).

 

Kinds of Niches:

No matter what concept or definition of niche is used, each niche can be measured in two ways:

 

Fundamental Niche:

Defines the maximum possible environmental limits of tolerance.

It is the maximum potential hypervolume of conditions, resources or roles that the organism could ever exploit.

Usually determined in laboratory experiments

Realized Niche:

Defines the actual limits of tolerance, resource use or roles that the organism actually experiences.

Usually determined in field studies.

 

 The Realized niche is almost always SMALLER than the Fundamental niche, for many reasons:

 

* Interactions between responses to different conditions (dimensions of the niche)

* Interactions between conditions and resources

* Interactions with "roles" of other organisms (competitors, predators, pathogens Ö)

* Habitat modification leads to adverse changes in the micro-environments actually experienced

* Historical reasons; time lags; the system is not in equilibrium, but the organism is expanding its ecological range

 

The Realized niche is sometimes LARGER than the Fundamental niche, for other reasons:

 

* Symbiotic associations with other species increase its tolerances, resource spectra or advantageous roles

* Habitat modification leads to favorable changes or buffering of micro-environments

* Historical reasons; time lags; temporary, non-equilibrial situation; contracting ecological range