Biology 20C - Fall 1998
ECOLOGY AND EVOLUTIONARY BIOLOGY
Lecture 1 - Introduction
Biological Themes Recurring in Ecology and Evolution: (Campbell: Chapter 1)
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Hierarchical organization of life |
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Emergent properties |
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Cell as a central unit of organisms |
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Individual as a central unit of ecology and evolution |
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Many scales (temporal, spatial, biological) |
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DNA provides continuity of life |
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Structure and function are closely related |
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Interacting themes of diversity and unity |
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Open systems of interactions with environment |
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Interacting processes of ecology and evolution |
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Scientific methods |
Ecology and Evolution:
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Both concerned with interactions between organisms and their environments |
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Both are intimately connected with processes of natural selection |
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Both emphasize processes within populations |
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Ecology "creates" the selective pressures that "drive" natural selection |
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Evolution "shapes" the ecological properties of organisms |
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They differ in their emphases on genetics |
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They often differ in their spatial and temporal scales. |
Aristotle (384-322 B.C.) can be viewed as the first "ecologist". Aristotle's ideas dominated western thinking about the natural world, and their widespread acceptance inhibited development of both ecological and evolutionary thinking for 2000 years. Four postulates in particular were not seriously challenged until the 18th, 19th and 20th centuries:
1. Eternal species |
Each species is fixed and never changes. [Therefore, change is NOT possible]. |
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2. Essentialism |
Each species has a fixed and "perfect" essence (idos). [Therefore, variation = "mistakes" in the expression of the idos, so variation cannot be important ( = "noise")]. |
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3. Scala Naturae |
(= Great Chain/Ladder of Being). The natural world is organized in a continuous gradation from small and simple to large and complex. It is a scale of increasing perfection leading to the Earth as the ultimate in physical perfection, over which man reigns as the ultimate in biological complexity and perfection. |
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4. Balance of Nature |
The natural world is in perpetual, "perfect" equilibrium. [Therefore, environmental and ecological variation is unimportant "noise" in the expression of this harmonious balance]. |
Acceptance of the Aristotle's first three postulates was sufficient to inhibit development of evolutionary thinking before the late 17th century, while acceptance of the fourth postulate effectively prevented development of modern ecological thinking before the 20th century. Belief in the balance of nature meant that even late 19th century ecologists were concerned mainly with static patterns, rather than with fundamental processes.
Aristotle's ideas became increasingly unsatisfactory beginning with the 15th century voyages of exploration that eventually forced Europeans to reconsider many ideas:
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Heliocentric rather than an Earth-centered solar system |
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New biotas shattered eurocentric views of homogeneous perfection and completeness |
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Geological evidence that the earth was very old |
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Fossils of extinct species raised questions of how it was possible for "perfect" species in a harmonious balance to become extinct |
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Multiple "creations" (>26) rather than a single biblical creation |
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Microscope gave more diversity and ideas of "spontaneous generation" of life |
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Modern taxonomy, from Linnaeus, challenged linearity of "scala naturae" |
By the late 17th century, many Aristotilean assumptions were being undermined, and European scholars were primed for a revolution in thinking. The final trigger was probably the Industrial Revolution, which demonstrated that, well within human lifespans, all aspects of the natural world could be altered drastically, without invoking divine intervention. A few key contributors were:
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Karl Linne (Linnaeus) (1753, 1758, 1767 +): Modern classification and taxonomy of living and fossil organisms. |
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Erasmus Darwin (Charles' grandfather): Expressed most of the components of natural selection, but as intellectual speculations, unsupported by evidence. |
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James Hutton (1785): Principle of Uniformitarianism |
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Jean Lamarck (1816 +): Non-genetic concepts of species change, evolution |
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Charles Lyell (1830-33): "Principles of Geology" |
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Charles Darwin (1829-32): Voyage of H.M.S. Beagle provided vast database |
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Charles Darwin (1859): Publication of "The Origin of Species" with evidence for natural selection |
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Alfred Wallace (1859): Working in southeast Asia had very similar ideas and evidence, independently of Darwin |
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Alfred Wallace (1876): "The Geographical Distribution of Animals" |
None of these people would have understood the distinction between what we call ecology and evolution today: they thought about both simultaneously. The fields were not clearly separated until after Louis Pasteur showed experimentally in the late 1800s that microbes did NOT generate spontaneously, and Gregor Mendel's work on inheritance became known at the turn of the 20th century, providing the basis of modern genetics. Nevertheless, Ernst Haeckel (1879) had already coined the term "oekologie" as a non-evolutionary concept, to describe studies of interactions between living organisms and their environments.
Biological Organizations
molecule - organelle - cell - tissue - organ - individual - population - community - ecosystem |
2. Dichotomous Hierarchies: (standard late 20th century view)
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tissue - organ - cell - organelle - molecule |
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[CELLULAR-MOLECULAR BIOLOGY] |
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individual |
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[ORGANISMAL BIOLOGY] |
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population - community - ecosystem - biosphere |
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community - ecosystem - biosphere |
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[ECOLOGY] |
molecule… cell… tissue… |
individual-population |
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[EVOLUTION] |
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species - genus…phylum…kingdom |
Ecological Scales:
Ecology is inherently interdisciplinary; it is concerned with syntheses among most branches of biology, as well as with many other sciences (e.g. geology, meteorology, oceanography, physics, chemistry…); it deals with complex questions spanning many spatial and temporal scales.
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Spatial |
Temporal |
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Ecological Levels |
Scales |
Scales |
(Relevant Disciplines) |
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Biosphere |
(whole earth) |
103 -105 km |
103 - 4.6x109 y |
(Biogeography, Evolution) |
Ecosystem |
(biotic + abiotic) |
101 -104 km |
102 - 106 y |
(Biol/Phys/Chem/Geol) |
Community |
(multiple species) |
100 -102 km |
101 - 103 y |
(Species interactions) |
Population |
(single species) |
100 -102 km |
100 - 103 y |
(Genetics, Demography) |
Individual |
(1 organism) |
10-6 -10-1km |
10-3- 102 y |
(Physiology, Behavior, Anatomy, Morphology) |
ECOLOGY (Campbell: Chap. 46)
Definitions:
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Oekologie is the study (logos) of organisms in their homes (oikos). (Haeckle, 1879) |
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Ecology is the scientific study of interactions between organisms and their environments. (Campbell, 1996) |
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Ecology is the scientific study of interactions affecting the distribution and abundance of organisms. (Andrewartha and Birch, 1954) |
Key Terms in Definitions of Ecology:
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Inductive |
"specific to general"; accumulate supporting evidence |
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Hypothetico-deductive |
"general to specific"; testable hypotheses; falsification |
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= " The Scientific Method" |
Both methods can use one or more of four approaches:
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Observational |
Mainly Field |
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Experimental |
Field and Laboratory |
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Theoretical |
Office |
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Modeling |
Office, Laboratory, Field (testing) |
B. Interactions: Modern ecology is concerned with understanding Processes rather than with describing Patterns. It deals with changes in space and time.
C. Organisms: Ecology is concerned mainly with whole organisms as individuals, and as groups of individuals (i.e. populations, communities and ecosystems)
D. Environment: In ecology, environment refers to all factors external to the organisms being studied:
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Biotic Environment |
Living organisms interacting with the target organisms, including food, competitors, predators, pathogens, parasites, symbionts … |
Originally the concept of environment was applied equally to individuals, populations, communities and sometimes even to ecosystems. The modern concept, dating from Andrewartha and Birch (1954), is that only an individual has an environment. This means that all other members of an individual’s own species are considered part of its environment. Andrewartha and Birch also introduced a classification of environmental factors under four general components. This classification has proven extremely useful for organizing ecological studies, and for defining field and experimental protocols. These components are:
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Weather. |
Most physical and chemical factors |
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Food. |
Usually other organisms, but includes nutrients for plants, and inorganic compounds for chemotrophic microorganisms |
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Other Organisms: |
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a. Different species |
Competitors, predators, pathogens, symbionts etc. |
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b. Same species |
Mates, family members, social groups, cannibals etc. |
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Place in which to live. |
Other requirements: nest sites, shelter… |
HISTORY OF ECOLOGY
Aristotle was recognizably an ecologist who studied nature solely by applying inductive reasoning to a strictly observational approach (e.g. he concluded from external morphology and seasonality that stalked barnacles were juvenile geese!). Because his postulates were the basis of ecological thinking for nearly 2500 years, they also delayed development of modern ecological thinking well into the 20th century. Belief in the balance of nature meant that ecologists tended to emphasize static patterns, rather than recognizing changes as fundamental ecological processes. Aristotlean postulates, especially the balance of nature, persist in popular thinking about ecology and the environment. The "Gaia Hypothesis" is perhaps it's most recent manifestation..
Plant and animal ecology developed along largely independent lines until the 1960s. Plant ecology developed relatively early in the 19th century, with strong biases to wards questions of distributions. Plants are large, easily seen, long-lived, and don't move (as adults). Their limits of distribution are readily detected and mapped. Plant ecology developed first as descriptive "plant sociology" before shifting its emphasis in the 20th century towards physiological studies concerned with identifying the physical or chemical factors limiting those distributions. Hence plant ecology emphasized low density populations in marginal habitats near the physiological limits of tolerance of the species.
Animal ecology developed towards the end of the 19th century, with its primary emphasis on questions of abundance rather than on distributions. This was reasonable, considering that most animals are very difficult to find or catch: they are usually small, highly mobile, often cryptic and secretive, and short lived. Since animals could be found easily only where they were abundant, animal ecology tended to concentrate on high density populations in favorable habitats. In most cases, animals' limits of distribution are unknown and very difficult to determine. By the 1920s, animal ecology was concerned with interactions among species (community ecology) and starting to recognize demography, the study of the structure and growth of populations (Elton, 1927)
Animal and plant ecology began to merge in the 1950s. Andrewartha and Birch (1954) merged the concepts of distribution and abundance by defining the limits of distribution as the place where the abundance falls to zero. In the 1960s, experimental studies of animal distributions became well established; and plant demography (concerned with the structure and abundance of plant populations) became an established discipline by the mid 1970s.
Like many other sciences, ecology diverged during the 20th century into many specialized fields (animal, plant, physiological, population, community, ecosystem, behavioral....ecologies) that tended to advance independently of the others. This trend has been reversed over the last 25 years, and the specialties are increasingly drawing on one another to form a synthesis in which each approach is treated more as looking at a different facet of the same phenomena. Similarly, the 20th century saw partitioning of evolution into many distinct disciplines (mendelian, quantitative, population, and molecular genetics; taxonomy and phylogenetics...) that are now coming together again in new syntheses. Finally, the late 19th century divergence between ecology and evolution, cast in concrete by the r spread of Mendelian genetics in the early 20th century, is also disappearing in such fields as "evolutionary ecology" or "ecological evolution".