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Microevolutionary Processes & HWE - Bio 1001 3.3, 3.4, 3.5 (p 48-58) 3.3 The Process of Evolution: How Does Natural Selection Work? • Lamarck and other researchers has already proposed evolution as a pattern in nature long before Darwin began his work • Darwin’s crucial insight lay in recognizing a process called natural selection Darwin’s Four Postulates • Darwin broke the process of evolution by natural selection into 4 logical sequences 1. The individual organisms that make up a population vary in the traits they posses, such as size and shape 2. Some of the trait differences are heritable, meaning that they are passed onto offspring genetically (height) 3. In each generation, many more offspring are produced than can possible survive. Thus only some individuals in a population live long enough to produce offspring and some will produce more than others 4. The subset of individuals that survive best and produce the most offspring is not a random sample of the population. Instead individuals with certain heritable traits are more likely to survive and reproduce • Natural selection occurs when individuals with certain characteristics produce more offspring than do individuals without those characteristics. The individuals are selected naturally by the environment • Evolution, which is any change in the genetic characteristics of a population over time, is simply the outcome of these 4 steps Biological Definitions of Fitness and Adaptation • Biological fitness; the ability of an individual to produce surviving offspring, relative to that ability in other individuals in the population • Fitness is a measurable quantity • The concept of fitness, in turn, provides a compact way of formally defining adaptation • In biology, an adaptation is a heritable trait that increases the fitness of an individual in a particular environment relative to individuals lacking the trait • Adaptations increase fitness - the ability to produce offspring Microevolutionary Processes & HWE - Bio 1001 3.3, 3.4, 3.5 (p 48-58) 3.4 Evolution in Action: Recent Research on Natural Selection Case Study 1: How did Mycobacterium tuberculosis Become Resistant to Antibiotics • Myobacterium tuberculosis, is the bacterium that causes TB • In Europe and the US, TB was once as great a public health issue as cancer is now • It receded in importance during the early 190s, for 2 reasons: 1. Advances in nutrition made people better able to fight off most M. tuberculosis infections quickly 2. The development of antibiotics such as rifampin allowed physicians to stop even advanced functions In the late 1980s, rates of the MTB infection surged in many countries, and in 1993 the • WHO declared TB a global health emergency • M. tuberculosis responsible for the increase were largely or completely resistant to rifampin and other antibiotics that were once extremely effective • How and why did the evolution of drug resistance occur? A Patient History • 2 months after the TB tests proved normal, however, the young man was readmitted to the hospital with a fever, severe cough, and laboured breathing • Despite being treated with a variety of antibiotics, including rifampin, he died of respiratory failure of 10 days later • Samples of material from his lungs showed that M. tuberculosis was again growing actively there • But this time, the bacterial cells were completely resistant to rifampin • Drug-resistant bacteria had killed the patient • Where did they come from? • After examining extensive stretches from each genome, the biologists were able to find only one difference: a point mutation in a gene called rpoB A mutation in Bacterial Gene confers Resistance • The rpoB gene codes for a component of the enzyme RNA polymerase • Recall, that RNA polymerase transcribes DNA to mRNA, and that a point mutation is a single base change in DNA • Rifampin, the drug that was being used to treat the patient, works by binding to the RNA polymerase of MTB • When the drug enters an MTB cell and binds to RNA polymerase, it interferes with transcription • If sufficient quantities of rifampin are present for long enough, and if the drugs bind tightly, bacterial cells don’t make proteins efficiently and they produce few offspring • But apparently the substitution of a leucine for a serine prevents rifampin from binding efficiently • Consequently, bacterial cells with the C --> T mutation continue to produce offspring efficiently, even in the presence of a drug Microevolutionary Processes & HWE - Bio 1001 3.3, 3.4, 3.5 (p 48-58) 1. Large population of MTB bacteria in patients lungs make him sick. By chance, a point mutation in the rpoB gene occurs in one or more bacteria cells 2. Drug therapy begins killing most MTB, patient seems cured and drug therapy is ended 3. The mutant cells proliferate, resulting in another major infection of the lungs. The patient becomes sick again 4. A second round of drug therapy begins but is ineffective on the drug resistant bacteria. The patient dies (pg 50) Testing Darwin’s Postulates Does the sequence of events illustrated mean that evolution by natural selection occurred? 1. Did variation exist in the population? Yes, Due to mutation, both resistant and nonresistant strains of TB were present prior to administration of the drug 2. Was this variation heritable? Yes. The researchers showed that the variation in the phenotypes of the two strains – from the drug susceptibility to drug resistance was due to variation in their genotypes 3. Was there variation in reproductive success? Yes. Only a tiny fraction of M.turbucolosis cells in the patient survived the first round of antibiotics long enough to reproduce 4. Did selection occur? Yes. When rifampin was present, certain cells – those with the drugresistant allele – had higher reproductive success than cells with the normal allele • • • • • M. tuberculosis individuals with the mutant rpoB gene had higher fitness in an environment where rifampin was present The mutant allele produces a protein that is an adaptation when the cell’s environment contains the antibiotic This study verified all four postulates and confirmed that evolution by natural selection had occurred The M. tuberculosis population evolved because the mutant rpoB allele increased in frequency Natural selection acts on individuals because individuals experience differential reproductive success o Only populations evolve o Allele frequencies change in populations, not in individuals When people hunt or fish, they may cause strong directional selection by harvesting the largest, oldest or fastest growing individuals in the population If mortality due to hunting or fishing exceeds mortality from natural causes, the result may be undesirable changes in the harvested population, as we select against the very traits we value Evolution by natural selection occurs when heritable variation leads to differential survival and reproductive success As a result, human predation swamps the selective effect of other causes of mortality Humans cause for more rapid phenotypic changes than other predators do Human harvested systems examined in the study showed phenotypic changes that were 300 percent higher than those in natural systems Microevolutionary Processes & HWE - Bio 1001 3.3, 3.4, 3.5 (p 48-58) Case study 2: Why are beak size, beak shape, and body size changing in Galapagos Finches? • • Peter and Rosemary Grnat (biologists) have been investigating changes in beak size, beak shape, and body size that have occurred in finches native to the Galapagos Island Early studies of the finch population established that beak size, beak shape, and body size vary among individuals, and that beak morphology and body size are heritable Selection during Drought Conditions • • • • • • • • • • • • In the annual season of 1977, Daphne Major received just 24 mm of rain instead of the 130 mm that normally falls The two observations support the hypothesis that most or all of the individuals died from starvation: o The researchers found a total of 38 dead birds, and all were emaciated o None of the missing individuals were spotted on nearby islands, and none reappeared once the drought had ended and food supplies returned to normal The research team realized that the die-off was a natural experiment o Instead of comparing groups created by direct manipulation under controlled conditions, natural experiments allow researchers to compare treatment groups created by an unplanned change in conditions o In this case, the Grants team could test whether natural selection occurred by comparing the population before and after the drought On average, survivors tended to have much deeper beaks than did the birds that died At the droughts peak, most seed sources were absent and thought the tough fruits served as the finches primary food source These fruits are so difficult to crack that they are ignored in years when food supplies are normal The group hypothesized that individuals with particularly large and deep beaks were most likely to crack these fruits enough to survive At this point, the Grants had shown that natural selection led to an increase in average beak depth in the population In only one generation, natural selection led to a measurable change in the characteristics of the population Alleles that led to the development of deep beaks had increased in frequency in the population Large, deep beaks were an adaptation for cracking large fruits and seeds The grants have documented continued evolution in response to continued changes in the environment Which genes are under selection? • Characteristics like beak size, beak shape, and overall body size are polygenic, meaning that many genes influence the trait (each one exerting a relatively small effect) Microevolutionary Processes & HWE - Bio 1001 3.3, 3.4, 3.5 (p 48-58) 3.5 – Common Misconceptions about Natural Selection and Adaptation • Research has shown that evolution by natural selection is often misunderstood • To help clarify how the process works, let’s consider some of the more common misconceptions about natural selection in light of data on drug resistance in the TB bacterium and changes in finch populations Selection Acts on Individuals • • • • Perhaps the most important point to clarify about natural selection is that during the process, individuals do not change – only the population does During the drought, the beaks of individual finches did not become deeper Rather, the average beak depth in the population increased over time, because deepbeaked individuals produced more offspring tan shallow beaked individuals did Natural selection acted on individuals, but the evolutionary change occurred in the characteristics of the population A contrast with “Lamarckian” Inheritance • • • • • There is a sharp contrast between evolution by natural selection and evolution by the inheritance of acquired characters Lamarck proposed that: 1) Individuals change in response to challenges posed by the environment 2) The changed traits are then passed on to the offspring In contrast, Darwin realized that individuals do not change when they are selected Instead, they simply produce more offspring than other individuals do Natural selection just sorts existing variants – it doesn’t change them Acclimation is not Adaptation • • Acclimation; is used to describe changes in individual’s phenotype that occur in response to changes in environmental conditions The key is to realize that phenotypic changes due to acclimation are not passed onto offspring, because no alleles have changed in composition Evolution is not Goal Directed • • • The mutation that conferred resistance did not occur because of the presence of the drug it just happened Every mutation is equally likely to occur in every environment. There is no mechanism that makes it possible for the environment to direct which mistakes DNA polymerase makes when it copies genes Adaptations do not occur because organisms want or need them its random Microevolutionary Processes & HWE - Bio 1001 3.3, 3.4, 3.5 (p 48-58) Evolution is not progressive; in fact complex traits are routinely lost or simplified over time as a result of evolution by natural selection There is no such thing as higher or lower organism • • • • • • • The non- progressive nature of evolution by natural selection contrasts sharply with Lamarck’s conception of the evolutionary process, in which organisms progress over time to higher and higher levels on a chain of being Under evolution by natural selection, there is no such thing as a higher or lower organism A human is no higher than its tapeworm parasite; each is well adapted to its environment All populations have evolved by natural selection based on their ability to gather resources and produce offspring All organisms are adapted to their environment, and are related by common ancestry Natural selection is not goal directed or progressive It simply favors individuals that happen to be better adapted to the environment existing at the time Organisms Do Not Act for the Good of the Species • • • Individuals with self-sacrificing alleles die and do not produce offspring. But individuals with selfish, cheater alleles survive and produce offspring Selfish alleles increase in frequency, while self-sacrificing alleles decrease in frequency Thus, it is not possible for traits that lead individuals to sacrifice themselves for the good of the species to evolve by natural selection Non-adaptive traits • • Vestigial traits are not the only types of structures with no function Some adult traits exist as holdovers from structures that appear early in development o For example; human males have rudimentary mammary glands o The structures are not adaptive o They exist only because nipples form in the human embryo before sex hormones begin directing the development of male organs instead of female organs Genetic Constraints • • • • • • Selection was not able to optimize all aspects of a trait In the case of the finches, wider beaks were not the best possible beak shape for individuals living in an arid habitat Wider beaks evolved anyway, due to a type of constraint called genetic correlation Genetic correlations occur because pleiotropy in which a single allele affects multiple traits Selection on alleles for one trait caused a correlated, though suboptimal, increase in another trait (beak width) Genetic correlations are not the only genetic constraint on adaptation Microevolutionary Processes & HWE - Bio 1001 • Lack of variation is also important 3.3, 3.4, 3.5 (p 48-58)