EVOLUTION

Essentials of  Biology Lecture

HOW POPULATIONS EVOLVE

HOW BIOLOGICAL DIVERSITY EVOLVES

 

 

Evolution provides the unifying concepts for the following chapters that deal with the diversity of organisms and the progressive development of derived anatomy and physiology. This section discusses the history and mechanisms of evolution, the history of life, and the most recent developments in classifying organisms.

Chapter 13/14 Darwin and Evolution

Discussion of Darwin’s formulation of a mechanism of evolution by natural selection is followed by description of the extensive and varied evidence for evolution.

Principles of Evolution  

The following is the first of our notes covering the principles of evolution.  Additional notes will follow.  

The modern theory of evolution explains the diversity of life from the perspective of science.  It provides a scientific explanation of how the multitude of species was created.  Advances in science, especially genetics, have strengthened this view of life in part by explaining the basis of variation and inheritance.  As you will see, variation and inheritance are both very important to the theory.  All aspects of modern biology are affected by the theory of evolution by means of natural selection.  It has been said that ‘nothing makes sense in biology except in light of the theory of evolution.’  Note the often used phrase, “The Darwinian Revolution”—a scientific revolution that has changed the focus of biology.  

Theories of Evolution  

I.  The Theory of Inheritance of Acquired Characteristics.  (1809) by  Jean-Baptiste Lamarck 

Modifications acquired during one’s lifetime are inherited by the next generation, ex. giraffes acquired a long neck slowly over time as each generation of giraffe stretched its neck slightly longer in trying to reach leaves high in trees.  At fist glance this theory is deceptively close to Darwin’s theory (both include the concept that evolution produces life forms adapted to their environments) but the inheritance of acquired characteristics implies that the organism itself can control the direction of change.  Unfortunately, there have been no discoveries of any such mechanism of change.  

II.  The Theory of Evolution by means of Natural Selection  

A.  The founders:

Charles Darwin  (1809-1882)  
English Scientist  
Age 22 - HMS Beagle  
Became ship's naturalist  
5 year voyage 1831-1836

Alfred Wallace (1823-1913)  
English Naturalist
1848-1852 Amazon R.
1854-1861 - Indonesia

Both men were excellent observers and prolific collectors and both worked as taxonomists.  
1858 - Wallace sends Darwin a letter, an essay in which he (Wallace) clearly communicates evolution by means of natural selection.  Darwin is known to have whined, "… all my originality [] will be smashed."  
1859 Darwin publishes On the Origin of Species by Means of Natural Selection  

B.  Seven Influential Factors for both Wallace and Darwin  

1).  Geology – study of the earth’s structure, origin, and history.  The book, Principles of Geology by Charles Lyell (1797-1875), includes the “theory of uniformity” or “uniformitarianism" - Geological processes have been uniform through time.  Geological changes of the past were caused by same, observable processes of today.  The process of change has been uniform through time.  Canyons and thick layers of sedimentary rock are the profound results of accumulated gradual change over vast stretches of time.  The significance to biology is this, slight changes over a long time have large-scale impact and one can look at the present to see the processes of the past.  Three concepts from geology are important in shaping the theory of evolution by means of natural selection:
·        the earth must be very old [4.6 billion years]  
·        one can look at the present to see processes of the past  
·        slight changes over a long time have large-scale impact  

2).  Fossils - extinct forms similar yet distinct from extant forms.  Perhaps a history of change (evolution) connects the living with the extinct.

3).  Island Life.  Isolated island populations differ slightly from nearby mainland populations.  e.g. Galapagos Islands (2 MYO, 600 mi. off W coast of S.A) the  birds, lizards, huge tortoises, similar to ‘nearby’ mainland species yet slightly different.  

Therefore there must have been Descent with modification (i.e., the island life evolved from mainland ancestors).  

4).  Overproduction of individuals - Thomas Malthus (British Economist) in his “Essay on the Principle of Populations”(1798) attempts to explain that much of human suffering, hunger, sickness, homelessness, and war are due to competition between an ever increasing number of individuals or groups of individuals for limited resources.  The number of individuals (population size) tends to increase exponentially.  This exponential capacity can’t be sustained (if it were, earth would be covered over many times with individual life forms).  Eventually, more individuals are born than can live to reproduce, leading to a struggle for existence.  Darwin and Wallace saw the concept relative to all species not just humans.  Only some individuals live to reproduce, many (in fact, most) will die before reaching reproductive age.  What determines, in nature [not human society] who lives and who dies?  An answer lies in part in variations among individuals.  

            [Social Darwinism is a distortion of the Theory of Evolution in the Darwinian sense, Social Darwinism was used by Hitler to justify the extirpation of Jews and his quest to destroy other countries; Antievolutionists use Social Darwinism as an attack on the Theory of Evolution.  Social Darwinism is the erroneous application of “survival of the fittest” to human societies.]  

5).  Individuals within a species vary extensively.  How does one know variations exist?  Taxonomy! A taxonomist observes and describes the variation within a species. You don’t have to be a taxonomist to know that individual people vary.  People vary not only in morphological traits such as hair texture and body shape but also in physiological traits as seen in the susceptibility of American Indians to European diseases [measles, mumps].  A knowledge of the variation in species of wild animal be they bird, beetle, or barracuda, requires the careful and detailed observation of many specimens.  Only a scrutinizing taxonomist, whether professional or amateur, sees the variation between individuals of wild species.  

6).  Many of the variations are inheritable and some variations may impart greater reproductive success, as in artificial selection below.  

7).  Artificial Selection – “selective breeding by humans of another species” (Harcourt Dictionary).  Variation among crops & livestock may be favored (selected) or disfavored (selected against), or neutral, i.e., neither favored nor disfavored, by man's control over which individuals are allowed to reproduce.  Man selects the best (according to his taste) and breeds these for future generations.  [see fig. 18.11-12, p. 290-291]

 

 

 

Charles Darwin and The Origin of Species

Charles Darwin's _______________________________________ was published on November 24, 1859.

Darwin argued that contemporary species arose from ancestors through a process of ____________________________ with ____________________ as the mechanism.

The basic idea of natural selection is that:

1. _______________________________________________

2. _______________________________________________

The result of natural selection is _________________________________________.

I. Darwin's Cultural and Scientific Context

The Origin of Species challenged the notion that the Earth was relatively _______________ and populated by _______________ species.

A. The Idea of Fixed Species

The Greek philosopher Aristotle held the belief that species are ____________ and do not ____________________.

The ___________________________ culture fortified this idea and suggested that the Earth is only 6000 years old.

B. Lamarck and Evolutionary Adaptations

In the mid-1700s, the study of ______________ began to form as a branch of science.

Naturalist Georges Buffon:

1. _________________________________________________

2. _________________________________________________

Jean-Baptiste Lamarck:

1. _________________________________________________

2. _________________________________________________

C. The Voyage of the Beagle

In December 1831, Darwin left _____________________ on the ______________________ to explore the world.

On his journey on the Beagle, Darwin:

1. _____________________________________________

2. _____________________________________________

Darwin was intrigued by:

1. _____________________________________________

2. _____________________________________________

D. The New Geology

Darwin was strongly influenced by the writings of geologist _______________________________

Darwin would apply Lyell's principle of _______________________________ to the evolution of Earth's life.

II. Descent with Modification

Darwin made two main points in The Origin of Species:

1. ____________________________________

2. ____________________________________

Evidence of Evolution

Biological evolution has left ________________________.

I. The Fossil Record

Fossils are preserved _________________ or _______________________ left by ___________________ that lived in the past.

They are often found in _____________________________.

The fossil record:

1. ____________________________________________________

2. ____________________________________________________

3. ____________________________________________________

____________________ are scientists that study fossils.  They have discovered many ___________________ forms that link past and present.

II. Biogeography

Biogeography is the study of ___________________________________.  Biogeography first suggested to Darwin that today's organisms evolved from___________________________.

Many examples of biogeography support _____________________________.

III. Comparative Anatomy

Comparative anatomy is the ________________________________________________.

Confirms that evolution is a _________________________________________________.

Homology is the _________________________________________________________.

_____________________________ are remnants of structures that served important functions in the organism's ancestors.

IV. Comparative Embryology

Comparative embryology is the study of _______________ that appear during the development of different organisms.

Comparative embryology of vertebrates supports ________________________________.

V. Molecular Biology

Evolutionary relationships among species:

1. ___________________________________________

2. ___________________________________________

Natural Selection

Darwin's __________________ are an excellent example of ____________________ and __________________________.

I. Darwin's Theory of Natural Selection

Darwin based his theory of natural selection on two key observations:

Observation # 1: _________________________

1. _________________________________________

2. _________________________________________

Observation # 2: ______________________________

1. _________________________________________

2. _________________________________________

Inference: _____________________________________________________________________

Those individuals with traits best suited for the local ______________________ leave more fertile offspring.

II. Natural Selection in Action

Examples of natural selection include:

1. ____________________________________

2. ____________________________________

3. ____________________________________

A. Does Predation Drive the Evolution of Lizard Horn Length?

Scientists concluded that horn length in these lizards was driven by ______________________.

Modern Synthesis: Darwinism Meets Genetics

The modern synthesis is the fusion of __________________ with ___________________________.

I. Populations as the Units of Evolution

A population is a group of individuals of the same _________________ living in the same _____________ at the same _______________.

A population is the smallest ________________________ that can evolve.

Population genetics focuses on populations as the ________________________ units.  It tracks the _____________________ of populations over time.

II. Genetic Variation in Populations

Individual ________________ abounds in populations.  Not all of this variation is ___________________.  Only the ____________________    of variation is relevant to natural selection.

A population is said to be _______________________ for a characteristic if two or more morphs, or forms, are present in noticeable numbers.

A. Sources of Genetic Variation

______________________ and ___________________ produce genetic variation.

Mutations: __________________________________________________

Sexual recombination: _________________________________________

III. Analyzing Gene Pools

The gene pool consists of all ____________________ of all individuals making up a population.

Alleles in a gene pool:

1. ______________________________________________________________

2. ______________________________________________________________

Genotypic frequencies:

1. ______________________________________________________________

2. ______________________________________________________________

The Hardy-Weinberg Formula:

1. ______________________________________________________________

2. ______________________________________________________________

II. Population Genetics and Health Science

The Hardy-Weinberg formula can be used to calculate the percentage of a ______________________ that carries the __________________ for a particular inherited disease.

III. Microevolution as Change in a Gene Pool

Hardy-Weinberg equilibrium describes a __________________________ population that is in genetic equilibrium.

Microevolution is defined as a ___________________________________ change in a population's frequencies of _______________.

Mechanisms of Microevolution

The main causes of microevolution are:

1. ___________________________

2. ___________________________

3. ___________________________

4. ___________________________

I. Genetic Drift

Genetic drift is a change in the gene pool of a small population due to _______________________.

A. The Bottleneck Effect

The bottleneck effect is an example of ____________IV. ___________.  Results from a ____________________ in population size.

Bottlenecking in a population usually reduces ___________________________.

B. The Founder Effect

The founder effect is genetic drift in a new ___________________.

C. Genetic Drift and Hereditary Disorders in Human Populations

The founder effect explains the relatively high frequency of certain ____________________ among populations.

II. Gene Flow

Gene flow is _____________________ with another population.  It tends to reduce ___________________ between populations.

III. Mutations

Mutations:

1. _________________________________________

2. _________________________________________

3. _________________________________________

IV. Natural Selection: A Closer Look

Of all causes of microevolution, only natural selection promotes ______________________.

A. Darwinian Fitness

Darwinian fitness is the contribution an individual makes to the ________________ of the next generation relative to the _________________ of other individuals.

B. Three General Outcomes of Natural Selection

Directional selection:

1. ___________________________________________

2. ___________________________________________

Disruptive selection:

1. ___________________________________________

Stabilizing selection:

1. ___________________________________________

Evolution Genetics of the Sickle-Cell Allele

Sickle-cell disease:

1. _____________________________________

2. _____________________________________

The sickle-cell allele confers resistance to the disease _____________________.

 

 

 

horizontal rule

 

Evidence of Evolution 

Before presenting some of the major lines of evidence for the fact that evolution has taken place, it might be useful to partition the scale of evolution into two categories:  1) the changes readily observed by population biologists today (microevolution), and 2) the profound changes which have accumulated from these population level processes (macroevolution).  Both scales of evolution provide evidence that evolution has occurred. 

Microevolution – the following three statements are meant to express the same basic concept and   define microevolution as: 1) change in the genetic makeup of a population, 2) change in a population’s allele or genotype frequencies, 3) change within the gene pool of a population.  Gene pool is simply a term created to express the concept of the collective total of all the genes in a population.  Basically, microevolution is evolution within a species such that gene frequencies change over time. 

Given an accumulation of population level changes, it is possible to look backward into history and see the larger scale of change, including the formation of new species, genera, families, etc. (macroevolution).  Evidence for microevolution comes from studies in population genetics.  Evidence for the larger scale of change is presented below.   

Macroevolution - origin of groups at and above the species level.  Macroevolution is said to be the “Grand View of Evolution” and includes the origin of new designs.  New designs are often reflected in the taxonomic hierarchy, e.g. the notochord is a new design unique to the phylum Chordata.  New designs are also reflected in phylogenies, e.g. a shared, derived character common to all deuterostomes is the new formation of the mouth from a 2nd embryonic derived opening into the animal body. 

Evidence of Macroevolution: 

1.  Fossils -- Paleontology- fossils are formed when sediment deposition buries dead bodies and the body remains undergo preservation via mineralization or an impression of body remains is preserved.  Fossils show: 1) change over time; & 2) continuity of change.

            ex. Archaeopteryx  - has teeth and long bony tail like reptiles but has feathers like birds.  Archaeopteryx is one of many known "missing links" between reptiles & birds.  

2.            Biogeography - study of the geographical distribution of plants and animals; it includes both the historical distributions as revealed by paleontology and present day distributions. 

            Mammal Distribution From Their origin 150 Million Years Ago To Today.  The concentration of diverse marsupial mammals in Australia (see p. 295) is thought to be the result of evolution within this group in the absence of placental mammals on this island continent.  Placental mammals, having evolved on northern continental masses, did not reach what is now Australia, thus leaving the marsupials to diversify without placental mammal competition.  (Bats are an exception, they are indigenous to Australia, other placental mammals did not reach the continent until the recent introductions by man—the rabbit, a placental mammal, is especially successful in Australia today as an exotic pest).  Presumably, placental mammals would have lead to the mass extinction of marsupial species.  At least in South America, the introduction of placental mammals from the north coincides with the extinction of many marsupial species known from the S. A. fossil record.  

3.  Comparative anatomy - study of body parts, esp. internal body parts.  The evolutionary view that all mammals share a common ancestor is supported by the anatomy of vertebrate forelimbs (see fig. 18.16).  Closely related species (e.g. species of mammals) show modification of shared structures, i.e., homologous structures. 

Homologous structures – “structures in different species that are similar because of common ancestry”; homologous structures have the same basic structure but may appear somewhat different depending on the degree of modification produced by evolution, ex. Whale flipper, bat wing, & human arm are all homologous (have same bones, are descended from common ancestor).  But a bat’s wing & a fly’s wing are analogous-similar in function but not origin or structure. 

Comparative anatomy reveals Vestigial organs – structures reduced in size & of little (or modified) function, e.g., in some snakes, such as pythons, evidence of their shared ancestry with legged creatures is found in their vestigial pelvic girdle and leg bones (for images see the following web link: Docent Webpage); some modern-day whales also have vestigial hind leg bones. 

4.  Comparative Embryology -  ex. Humans, like all vertebrates, have a notochord, post anal tail and gill pouches as embryos (see p. 296 fig. 18.17).  Such evidence does support the view that all vertebrates share a common ancestor and that the differences between vertebrates today is the result of “descent with modification” from this ancestral type.  A replay of the changes from the ancestral type was once thought to exist in the changes during embryonic development.  The well-worn phrase "ontogeny recapitulates phylogeny" (or stated more clearly, embryonic development recapitulates evolutionary history) is a gross overstatement.  The point is, that useful features to unraveling the evolutionary relationships among animals can be learned from comparative embryology. 

5.  Molecular Biology – The genetic code is universal.  Also, degrees of relatedness are reflected in nucleotide base sequence (DNA) similarity and in amino acid sequence similarity in proteins.   For example, horse hemoglobin amino acid sequence is more similar to the sequence in hedgehog hemoglobin (both the horse and hedgehog are mammals) than to the sequence in hummingbird hemoglobin (see fig. 18.18 for similar example).

horizontal rule

 

 

Principles of Evolution  

Natural Selection – the mechanism of evolution as told by Wallace and Darwin

**Natural Selection - differential reproductive success that results from the interaction of organisms with their environment.** 

I.  Some Background 

Remember three important subtle points:  1)  Natural selection can amplify or diminish only heritable variations.  Only heritable variations are affected; acquired characteristics play no role in evolution.2)  Populations evolve, not individuals.3)  Natural selection is situational (the direction it takes depends on the environment, i.e., the selective forces). 

To illustrate these subtle points using the case of the peppered moth… 1) moth color is inherited perhaps under control of a single gene; 2) the changes in nature over time involve whole populations, that is, in pre-industrial times the dark moth form was quite rare and perhaps in the distant past the dark mutation had not yet appeared.  With time and natural selection at work, the dark moth became the most common form within the population.  3) the 3rd subtle point is quite important, for what is advantageous in one environment may not be in another.  In the saga of the peppered moth, under polluted tree bark conditions following the industrial revolution the dark from became most common, yet a return to a predominance of light colored forms follows restoration of healthy, non-polluted tree bark.  (The story of the peppered moth has additional complexities but we will ignore them). 

A. About heritable variations
Heritable variations are governed by genes, and as we have seen, genes are mere chemicals containing coded information in the form of DNA.  DNA’s ability to mutate is the fundamental basis for evolutionary change.  While this primary source of variation is the result of random, chemical events (i.e. gene mutations), other aspects of the process of evolution (e.g. natural selection) are very nonrandom, as we will see.    

B.  About Populations
Natural Selection (evolution) is a population level phenomenon, i.e. populations evolve, not individuals, and furthermore, the variations upon which natural selection operates exist within populations (variations exist only in the context of populations). 

Population - a) a group of individuals of the same species, b) live together in same area at same time, c) exchange genes (reproduce sexually).

Sources of Variation w/in populations:
     1) Mutation - creates new alleles/new traits/new phenotypes- The fundamental basis of change
     2) Sexual reproduction - recombines alleles (recall Punnett-squares)
          Benefit of sexual reproduction: genetic variation in offspring allows some to survive in changing environment (e.g. clonal algae turn sexual
         during unfavorable conditions)
     3) Gene Flow - gain or loss of alleles (genes) from a population  
          gain alleles - immigration (arrival of new individuals)    lose alleles - emigration or death 

C.  About Situations or Circumstances
The type of change natural selection can bring about depends on the situation or circumstances of the environment.  For example, for a population of domesticated, pedigree dogs the continued existence of each breed is dependent on an environment in which owners continue to breed pure lines for the survival of the breed.  Perhaps of more interest is the situation (environment) in which breeders desire new combination of characters and experiment with hybridization between breeds.  Breeders hope to develop new lines of pure breeding dogs in this way.  Each decade, new breeds of dogs, cats, rabbits, etc. are created in this process (=artificial selection) that is not unlike the process of natural selection.   

Expounding on the situational aspect in which evolution operates, Stephen Jay Gould, Harvard paleontologist and prolific writer popularizing evolutionary concepts, puts fourth an interesting idea that he calls the “contingency of evolutionary history” in his 1989 book, Wonderful Life: The Burgess Shale and the Nature of History.  The title was inspired by the 1946 movie It’s a Wonderful Life in which George Bailey learns how his own life was a contingency for many other events.  One such contingency that became a prerequisite for human evolution was the extinction of the dinosaurs that made way for mammal diversification.  Apparently, dinosaur extinction was contingent on an asteroid that impacted Earth 65 million years ago.  Had these events not occurred, humans may never have evolved.  (The question of why humans evolved (to what purpose) is not a question science can address; the question of how humans evolved is explored through the process of science.  See chapter 21 if you are interested in human evolution.) 

II.  Types and examples of Natural Selection  

Genetic variation within a population creates, in part, a diversity of phenotypes, phenotypes that have a strong genetic component (as opposed to environmentally induced phenotypic differences).  Depending upon the situation/environment or circumstances affecting survival, certain variant phenotypes may enjoy greater reproductive success.  Those aspects of the phenotype that correlate with greater success we think of as adaptations, e.g. the shell of reptilian eggs is an adaptation to life on land.  Adaptations result from natural selection.  

The following examples illustrate differential reproductive success (i.e., natural selection) as driven by different scenarios of selection pressures. 

1.  Stabilizing selection - favors intermediate phenotype
                          - does not lead to change
                          - extremes selected against
                          ex. Birth weight in humans  and   gall fly gall size on golden rod

2.  Directional selection - important when environmental change occurs
            ex. resistance to pesticides/antibiotics
                        - favors an extreme phenotype, leads to changes in populations & eventually (if you accept theory of uniformity, uniformitarianism)                           to new species formation.

The origin of the hard, reptilian shelled egg can also be thought of in terms of directional selection coupled with genetic mutations leading to ever more protective egg coverings evolving from jelly-like frog egg-ish beginnings.  Selection pressures would favor those eggs that survive greater desiccation as experienced in the ever more terrestrial lifestyles of the early reptiles.

3.  Disruptive (Diversifying) Selection - the intermediate phenotype selected against.
            ex. predator favors medium sized prey

In all 3 modes: 
Survival of the Fittest where fitness=reproductive success (not strength)

4.  Sexual selection - Selection for mate attracting traits
                                  ex. Male’s bright plumage in birds, males greater strength, larger antlers

Sexual selection provides an explanation for sexual dimorphism - male & female differ  beyond gonads and genitalia.  Recall the extreme case of the dwarf male angler fish.  Based on a morphological species concept alone, male angler fish may have been named as a separate species when originally captured from the depths of the ocean.  Only after the realization that the small angler fish were simply  males which spawned with the much larger female angler fish would a single species rather than two be recognized.  The knowledge of the capacity for interbreeding is of course the basis for the biological species concept.

 

 

 

Back to the top