5.

4 EVOLUTION (8) (Kent chapter 20 page 436) The following is taken from Kimballs Biology Pages an online resource.

Evolution and Adaptation

In 1859 the English naturalist Charles Darwin published The Origin of Species. The book contained two major arguments: First, Darwin presented a wealth of evidence of evolution. He said that all living things on earth today are the descendants with modifications of earlier species. Second, he proposed a mechanism natural selection to explain how evolution takes place. Evolution involves two interrelated phenomena:
o

adaptation Over the course of time, species modify their phenotypes in ways that permit them to succeed in their environment. This page is devoted to looking at how evolution leads to adaptation.

speciation Over the course of time, the number of species increases; that is, a single species can give rise to two or more descendant species. In fact, Darwin maintained that all species are related; that is, any two species on earth today have shared a common ancestor at some point in their history.

THE ASSESSMENT STATEMENTS 1. Define evolution. Evolution is the cumulative change to the inheritable characteristics of a population. Gene pools of species change. New species form from pre-existing ones. All life is connected due to its common origins. 2. Outline the evidence for evolution provided by the fossil record, selective breeding of domesticated animals and homologous structures. FOSSIL RECORD (p438) Fossils indicate that a large range of organisms have lived on earth in the past that are morphologically different to todays living species. Modern species are not found as fossils mixed in with the different ancient forms.

Problems: The fossil record provides too little data to be able to trace continual change for species. Evidence is widely distributed geographically, often concentrated in locations that favoured the fossilising process. The dating processes provide an approximate age only. Soft bodied organisms do not fossilise easily and are scarce in the fossil record. SELECTIVE BREEDING OF DOMESTICATED ANIMALS (p448) Animal breeders mate animals with characteristics they wish to see in the progeny. Through a continued process across many generations this selection creates a reproducible set of characteristics in the population that we call a breed. The gene pools of each breed contain a unique common set of alleles. Dogs of many breeds are an example of this. Todays breeds of dogs have evolved from a pre-existing population of dogs under the control of humans. This is an artificial form of selection as opposed to natural selection. Humans have used the process of natural selection to change dogs in a way that suits them. Is this morally justifiable? HOMOLOGOUS STRUCTURES (p441) Structures found in different species that have a common pattern or organisation. A classic example is the pentadactyl limb of mammals. (p 441) It is a skeletal pattern found in arms, forelegs, fins and wings of mammal species adapted to surviving in different environments. They all contain similar ones in the same relative positions. MECHANISMS FOR EVOLUTION (20.4 p 442 Natural Selection) 3. State that populations tend to produce more offspring than the environment can support. Either through rapid breeding rates or multiple offspring, when combined with lifespan most organisms can produce many more offspring organisms than is needed to simply replace themselves. The effect of this is a tendency to increase both the population and the competition within the population for the resources needed to sustain that population. 4. Explain that the consequence of the potential overproduction of offspring is a struggle for survival. (p442) As the population attempts to survive there is an ongoing struggle with the environment such that even in isolation a rapidly increasing population becomes self limiting as resources per capita diminish. (food, shelter, etc) Competition between members of a species creates a struggle for survival that results in a tendency for the survival of individuals better able to survive where others of the species die or are unable to reproduce. 5. State that the members of a species show variation. Individuals of a species are not identical physically or physiologically. They do share a lot of genes but subtle differences between them are due to each having a unique set of alleles. Thus, within all species the individuals show variation in structure and

function to each other. These variations are so subtle however, that the individuals look and function in a very similar way and can be identified as a separate but unified group of organisms. 6. Explain how sexual reproduction promotes variation in a species. Sexual reproduction involves the process of meiosis for the production of gametes. This creates variation in individual gametes formed due to: a) random alignment of chromosomes at metaphase 1 leads to independent assortment. This means that in sexual reproduction, the same parent organisms never produce identical offspring from separate mating events. b) Formation of chiasma and the resulting crossing over of alleles between non sister chromatids during stage 1 c) There is a chance for genetic mutation to occur during DNA replication that happens for the formation of gametes. and d) The random process by which two gametes come together for fertilisation leads to variations in allele associations that result in a different mix of phenotypes. 7. Explain how natural selection leads to evolution.

Natural Selection

Living things produce more offspring than the finite resources available to them can support. Thus living things face a constant struggle for existence. The individuals in a population vary in their phenotypes. Some of this variation is inheritable; that is, it is a reflection of variations in genotype. Those variants best adapted to the conditions of their life are most likely to survive and reproduce themselves ("survival of the fittest"). To the extent that their adaptations are inheritable, they will be passed on to their offspring.

The forces of natural selection act on phenotypes, but only if there is a change in the genotypes of a population has evolution occurred. Extra information- again it is from Kimballs Biology Pages.

The Measure of "Fitness"

Fitness is a measure of reproductive success. Those individuals who leave the largest number of mature offspring are the fittest. This can be achieved in several ways:

Survival or mortality selection Mating success or sexual selection Family size or fecundity selection

Survival
Any trait that promotes survival at least until one's reproductive years are over increases fitness. Such traits are adaptations.

Sexual Selection
In sexual selection, one sex usually the female chooses among the available males. Any inherited trait that improves the mating success of certain individuals will become more pronounced in succeeding generations. Some examples: When ready to mate, female three-spined sticklebacks (fish) choose males with many Class II MHC alleles over males with fewer alleles. Class II alleles encode the proteins that present antigens to the immune system. Presumably, the more of them you have, the greater the diversity of parasite antigens your immune system can recognize and defend against. Link to discussion of how polymorphism of MHC alleles protects against parasites. The females distinguish between the males by soluble molecules ("odors") the males release into the water. How these "odors" are controlled by the MHC alleles is not known. A culture of Drosophila set up with equal numbers of red-eyed and white-eyed flies of both sexes will, after 25 generations or so, end up having only redeyed (the "normal") flies in it. This, despite the fact that white-eyed flies are just as healthy and live just as long as red-eyed flies, i.e., they are equal in terms of survival. But, as it turns out, not only do red-eyed females prefer redeyed males, but white-eyed females do also.

In other cases of sexual selection, one phenotype prefers to mate with others of the same phenotype. This is called assortative mating.

Fecundity Selection
The production of a large number of mature offspring is a measure of fitness. I stress mature because only they can pass these traits on to another generation. Some ways to do this:

Earlier breeding. If some females become sexually mature earlier than others, their chances of leaving offspring are enhanced. For some species (e.g., fish, oysters), which provide little or no care for their young, fitness is measured by the number of fertilized eggs they produce. For species (such as ourselves) that take care of their young; selection acts to reduce family size (to a point). A large study in Utah (U.S.A) showed that in the 19th century families with fewer children had more surviving grandchildren.

8. Explain two examples of evolution in response to environmental change; one must be antibiotic resistance in bacteria. (I suggest the other to be prevalence of sickle cell allele in conjunction with malaria as it reinforces other core course content. See spread 16.1 page 341 AB ) Ruw ANTIBIOTIC RESISTANCE (The information below should be studied and summarised)

Resistance to Antibiotics
None of the antibiotics discussed below is effective against all bacterial pathogens.

Intrinsic resistance
Some bacteria are intrinsically resistant to certain of the antibiotics. Example: Grampositive bacteria are much less susceptible to polymixins than Gram-negative bacteria. [The "Gram" designations refer to the reaction of the bacteria when stained with the Gram stain; this reaction is a reflection of the very different organization/stucture of their cell walls.]

Acquired resistance
Many bacterial species acquire resistance to one or more of the antibiotics to which they were formerly susceptible. Example: In the U.S. in the decade from 19851995, resistance of Shigella (which causes gastrointestinal illness) to ampicillin grew from 32% to 67%. And, while only 7% of these isolates were resistant to the combination of sulfamethoxazole and trimethoprim at the start of the decade, that figure had grown to 35% by the end of the decade. Bacteria develop resistance by acquiring genes encoding proteins that protect them from the effects of the antibiotic. In some cases the genes arise by mutation; in others, they are acquired from other bacteria that are already resistant to the antibiotic. The genes are often found on plasmids which spread easily from one bacterium to another even from one species of bacterium to another. Examples:

Synthesis of the enzyme penicillinase or other beta-lactamases provides protection from the beta-lactam antibiotics. These enzymes break the beta-lactam ring at the position shown with the green arrow in the diagram of penicillin G. Likewise synthesis of cephalosporinases defeats the cephalosporins. Defeating quinolones: o Some bacteria do this by modifying their DNA gyrase. o Others, e.g., Mycobacterium tuberculosis, develop quinolone resistance by synthesizing a protein that resembles a short length of DNA. This protein binds the gyrase so it cannot form the DNA/gyrase complex that is the target of quinolone action.

Some bacteria synthesize "pumps" in their plasma membrane through which they remove antibiotics like tetracyclines from the interior of the cell. Bacteria may methylate their ribosomes obscuring the target of antibiotics (e.g., erythromycin) that ordinarily bind to and inactivate the ribosome or conversely they may enzymatically modify the antibiotic (e.g., kanamycin) so it can no longer "see" its ribosomal target. Bacteria may modify the structure of their peptidoglycan wall and thus avoid the inhibitory effects of antibiotics like cycloserine.

An alarming number of human pathogens have acquired genes to combat all the presently-used antibiotics except vancomycin and recently vancomycin-resistant bacteria have appeared. These multidrug-resistant strains are particularly common in hospitals where antibiotic use is heavy, and the patients often have weakened immune systems. How transposons contribute to multidrug resistance.

Measuring Antibiotic Resistance

The figure illustrates the simplest method of the several available for measuring antibiotic resistance.

A suspension of the bacteria to be tested (e.g. cultured from the infected patient) is spread over the surface of a petri dish containing a solid culture medium. Disks of several different antibiotics are pressed on the surface of the agar. The concentration of antibiotic in each type of disk is standardized. Incubate overnight. The bacteria will grow into a "lawn" except where an antibiotic to which they are sensitive has diffused out from its disk. Measure the diameter of any zones of inhibition that are formed.

What can you do to delay the spread of antibiotic resistance?

Don't ask your doctor for an antibiotic to treat a viral disease (e.g., a cold) for which antibiotics are useless. (However, your doctor may prescribe an antibiotic if you are infected by an influenza virus not to fight the virus but to protect you against a secondary bacterial infection of your damaged lungs.) Stay the course. Use all doses prescribed even though you are feeling better. This will minimize the opportunity to select for resistance among the bacteria that remain late in the infection. Don't save unused antibiotics for later self-medication.

Farmers can help as well by avoiding the use of antibiotics in their livestock that are similar to those used in humans. Antibiotics are widely used in healthy livestock to improve their growth rate (by an unknown mechanism).

An article in the 20 May 1999 issue of The New England Journal of Medicine documents the recent development of quinolone resistance in Campylobacter jejuni, the most frequent bacterial cause of gastroenteritis in humans. The rise coincides with the approval in 1995 of the use of quinolones by U. S. poultry farmers (chickens also become infected by C. jejuni). Similar recent increases in fluoroquinolone-resistant C. jejuni have been reported in the Netherlands and also in Spain (where as many as 50% of human infections are now caused by bacteria resistant to the antibiotic). In each country, the appearance of resistant strains followed the widespread introduction of quinolone treatment for animals.

Future Prospects
Drug companies after many years of complacency are now responding to the threat of antibiotic-resistant bacteria. Over a dozen new antibiotics are being developed and some have already reached clinical trials. Many of these are semi-synthetic modifications of already-existing antibiotics, including new

beta-lactams macrolides glycopeptides quinolones modifications of vancomycin

Others are entirely new, attacking previously-unexploited chinks in the bacterial armor.

Urea hydroxamates, that block the enzyme (peptide deformylase) that removes fMet from the finished protein so it can begin its work. (Eukaryotes do not begin translation with fMET.) Heteroaromatic polycycles (HARP) that bind to bacterial promoters preventing gene transcription.

Summary Any population of bacteria contains individuals with a range of phenotypes. When an antibiotic substance is applied to such a population it is probable that some individuals will not be affected by it. The dead or damaged bacteria cannot reproduce and no longer play a part in the evolution of that species. The surviving bacteria reproduce to create a new population with a higher proportion being unaffected by the antibiotic substance. When the same substance is used in an attempt to control another infection the proportion of surviving bacteria is greater than before. In this way the species becomes resistant to the antibiotic effects of the substance to such a level that it no longer is useful in treating disease caused by it. The population has undergone a change in its genome; this is evolution by natural selection.

Of course bacterial species are not confined and the application of one antibiotic to one bacterial population does not mean the entire population of that bacterium worldwide has changed. This is how various strains of bacteria come to exist within the same species. This is similar to breeding programmes that produce different breeds/varieties of animals and plants. Balanced Polymorphism an example of natural selection altering the proportion of genes in a population. In regions of the world (e.g., parts of Africa) where malaria caused by Plasmodium falciparum is common, the allele for sickle-cell haemoglobin is also common. This is because children who inherit

one gene for the "normal" beta chain of haemoglobin and one sickle gene

are more likely to survive that either homozygote. Conversely,

Children homozygous for the sickle allele die young from sickle-cell disease but children homozygous for the "normal" beta chain are far more susceptible to illness and death from falciparum malaria than are heterozygotes.

Hence the relatively high frequency of the allele in malarial regions. Here is another example of natural selection, this time relating to heavy metal tolerance in plants.

Disruptive Selection
In some circumstances, individuals at both extremes of a range of phenotypes are favoured over those in the middle. This is called disruptive selection. An example: The residues ("tailings") of mines often contain such high concentrations of toxic metals (e.g., copper, lead) that most plants are unable to grow on them. However, some hardy species (e.g. certain grasses) are able to spread from the surrounding uncontaminated soil onto such waste heaps. These plants develop resistance to the toxic metals while their ability to grow on uncontaminated soil decreases. Because grasses are wind pollinated, breeding between the resistant and non-resistant populations goes on. But evidently, disruptive selection is at work. Higher death rates of both

less resistant plants growing on contaminated soil and more resistant plants growing on uncontaminated soil

leads to increasing divergence of the populations into two subpopulations with the extreme manifestations of this trait.

The evolutionary significance of disruptive selection lies in the possibility that the gene pool may become split into two distinct gene pools. This may be a way in which new species are formed. The formation of one or more species from a single precursor species is called speciation. This is evolution!