Biology 1M03 Complete Review

Patrick Te Biology 1M03, 2011
Biology 1M03 Review

Worst course ever but yeah
Biological Science
Chapter 1: Biology and the Tree of Life Biological science was founded partially with the development of two things o The cell theory. o The theory of evolution by natural selection. Phylogenetic trees are diagrams that may be interpreted as a geographical representation of inferred evolutionary relationships among species. o Phylogenies can be established through analyzing similarities/differences in traits. Biologists follow the same processes all scientists do. o They ask questions, generate hypotheses and design experiments to see if they can deny or support a hypothesis. o A typical scientific theory has two parts. Identification of a pattern. Identification of a mechanism or process responsible for this pattern. The Cell Theory 1660s: Hooke, van Leeuwenhoek first scientists to observe cells. o Cells are highly organized compartments bound by a plasma membrane containing concentrated chemicals in an aqueous solution. o The cell theory states that all organisms are made of cells and all cells come from preexisting cells. Louis Pasteur: Demonstration that cells come from other cells and are not generated spontaneously. o Two nutrient broths, one with a straight neck flask, the other with a swan neck. o Straight one had cells in it later on, swan one did not (they could not enter), therefore cells were deemed to not be spontaneously generated. Assuming all cells really do come from preexisting cells, this implies that all individuals in a population of single celled organisms would have to be related by common ancestry. The same idea applies to multicellular organisms. The Theory of Evolution 1858: Darwin and Wallace proposed that all species are related by common ancestry. o Also proposed that the characteristics of species can be modified from generation to generation. Evolution entails that species are related to one another and can change over time. Natural selection is a process that explains how evolution occurs. Fitness may be conceptualized as the ability of an individual to survive and produce offspring. Adaptation ( on a microscopic scale ) may refer to a trait that increases the fitness of an individual in a given environment.
Natural Selection and Populations A group of individuals of the same species living in the same area constitutes a population. Two conditions must be met for natural selection to occur: o Individuals in a population vary among themselves in heritable characteristics. o In a given environment, certain versions of these heritable traits help individuals survive better or reproduce more than other versions of the traits. When certain heritable traits lead to increased success in surviving / producing offspring, then these traits will become more common in the population. Natural selection acts on individuals, but evolutionary change affects populations. Evolution occurs when heritable variation leads to differential success in reproduction. Artificial selection defines individuals in a population being selected for mating based on particular traits. Repetition of this process over generations changes the characteristics of a domesticated population over time. The Tree of Life The cell theory and theory of evolution imply that all species come from preexisting species and that all species past and present can trace ancestry back to a single common ancestor. Spectiation is a divergence process in which natural selection has caused populations of one species to diverge, forming new species. The tree of life is a diagram that may be interpreted to infer genealogical relationships among species with a single ancestral species at its base. Carl Woese and co. o Studied small subunit rRNA as a means of understanding the evolutionary relationships among organisms. rRNA sequences should be very similar in closely related organisms and less similar the less related organisms are. A phylogenetic tree is a diagram that can be seen as depicting these relationships among species. Branches close to one another represent closely relates species, while farther apart branches indicate more distant relation. The tree of life indicates 3 main groups of organisms, the eukaryotes and two groups of prokaryotes (bacteria and archaea) o Creation of a new taxonomic level called the domain, consisting of bacteria archaea and eukarya. The tree continues to change as the location of many branches remains debated. Taxonomic Levels Created by Linnaeus, it is a hierarchical system, going from least to most specific. 1. Kingdom 2. Phylum 3. Class 4. Order 5. Family 6. Genus 7. Species
Patrick Te Biology 1M03, 2011 In terms of kingdoms, Linnaeus proposed two kingdoms of plants and animals. An alternate fivekingdom system was later proposed.
Chapter 24: Evolution by Natural Selection Populations and species evolve. o Allele frequencies, and by extension, characteristics of species change over time. Natural selection occurs when individuals with certain alleles survive or produce the most offspring in a population. o Adaptations on a microevolutionary scale are genetically based traits that increase ability to survive or produce offspring in a given environment. Evolution by natural selection is not progressive and changes characteristics of the population, not individuals. o Animals rarely do things for the good of the species (selfish alleles being more common), except under particular conditions (ex. Protection of young). Introduction Scientific theories consist of observations about a natural pattern and a proposed process explaining the pattern. Charles Darwin and Alfred Russell Wallace 1858 o Original people that proposed evolution, that is, the idea that species on earth have changed over time. They explained the pattern of evolution through the process of natural selection. o Evolution is currently one of the most supported scientific theories in the history of research. The Evolution of Evolutionary Thought Plato o Proposed that every organism was an example of perfect essence or type created by God. These types were unchanging. o Theory of special creating = typological thinking = variations are unimportant. Aristotle o Proposed the ordering of organisms into a great chain of being. All species were organized in a sequence that focused on the size and complexity of the organism (humans topped this chain). o Ex.) Unicellular organisms -> fungi -> green algae -> land plants, etc. Jean-Baptiste de Lamarck (1809) o First to propose a formal theory of evolution. o Simple organisms start at the base of the great chain though spontaneous generation. Evolution takes place through moving up the chain over time and is a linear pattern. Escalator analogy instead of the ladder. o Inheritance of acquired characteristics was responsible for this pattern of change. Individuals change in response to an environment, passing on these changes to offspring. (Single-generation change) Darwin and Wallace o Proposed that change in species over time is NOT linear, instead being based on variation among individuals in populations through natural selection.
Patrick Te Biology 1M03, 2011 A population is defined as individuals of the same species living in an area. Natural selection is the process by which individuals of a population with certain heritable traits tend to survive and produce more offspring compared to individuals without these traits. This eventually leads to a change in the makeup of the population.
The Pattern of Evolution Darwin described evolution as descent with modification; change over time means that modern species originated from ancestral species. Therefore o Species change over time. o Species are related by common ancestry. Evidence for Change through Time Fossils are traces of past organisms. The many found and described fossils make up our fossil record. Most fossils are found in sedimentary rock, formed from layers of sand or mud. o Layers of sedimentary rock are associated with different intervals in a geologic time scale, a relative scale based upon fossil content. Time divided into eons, eras, period and epochs. Researchers now use radioactive isotopes (carbon dating) to assign actual years to the geological data. o The Earth is approximately 4.6 billion years old, with the earliest discovered signs of life having existed 3.4 billion years ago. o Fossils provide evidence for extinct species. Darwin interpreted extinction as supporting his theory. If species have indeed gone extinct, it means that the species living on earth have changed over time. Extinct fossil species were typically succeeded by similar species. (Law of Succession). Darwin interpreted this as evidence that supported the notion that extinct and living forms are related (ancestors, descendants) Transitional Forms: Traits between both earlier and later species exhibited. Vestigial Traits Defined as traits that have been reduced or incompletely developed, resulting in either reduced or no function. They are, however, clearly similar to functioning structures found in related species. o Additional evidence that characteristics of species have changed over time. Ex.) Human coccyx (tails, monkeys), goosebumps (erect body hair, chimps) Geographic Relationships Darwin collected mockingbirds from the Galapagos Islands that, while superficially similar, varied in species from island to island. o Proposed that the similarities were due to the fact that they had descended from a common ancestor, evolving differently depending on location to survive. o Related in a phylogeny. Relationship shown on a phylogenetic tree. Genetic and Developmental Homology
Patrick Te Biology 1M03, 2011 Homology is a similarity that exists between species that have descended from common ancestry. Homology is recognized on three interacting levels. o Genetic Homology: Similarity in the DNA sequences of different species. (Genetic code) o Developmental Homology: Similarity in embryonic traits. (Gill pouches found in chicks, humans, and cat embryo) o Structural Homology: Similarity in adult traits. In many cases, traits are similar between species because the species in question are related by common descent. The likelihood of these similarities occurring is less if they were created independently of one another. Ex.) Similarities in hand structure between humans, horses, birds and turtles.
How does Natural Selection Work? Darwins Four Steps 1. Individuals that make up a population vary in their traits. (Variation) 2. Certain traits are heritable, that is capable of being passed to offspring. (Heritability) 3. In each generation, many offspring are produced, but not all survive. Only some individuals survive and produce offspring. (Competition) 4. Individuals with certain heritable traits are more likely to survive and reproduce. Natural selection occurs when individuals with certain traits produce more offspring than individuals that do not have them. (Skewness) a. These traits will increase in frequency from one generation to the next. Evolution results because of this, as genetic characteristics change with the physical characteristics of a population overtime. Modern biologists have condensed Darwins process into two statements. Evolution by natural selection occurs when... 1. Heritable variation leads to 2. Differential success in survival and reproduction. Evolution can be more specifically defined as a change in allele frequencies in a population over time. Example: Depending on their genotype, a certain kind of moth is black, grey or white. If birds eat the black moths more than the white ones, then, naturally, the white moths will be able to survive longer and produce offspring. As this trend continues, white moths eventually become more common than black ones in the moth population. The Process of Evolution: Fitness and Adaptation Darwinian fitness defines the ability of an individual to survive and produce offspring relative to the ability of other individuals in a population. In biology, an adaptation on the micro evolutionary scale is a heritable trait that increases Darwinian fitness in a particular environment relative to individuals lacking that trait. Examples of Natural Selection M. tuberculosis o Disease that killed many people in the 19th century (up to 1/3rd of all adults in cities). o Sanitation, nutrition and antibiotics greatly reduced the number of cases by the 1950s up to around 1990. o Rates of TB surged again in 1980 due to the evolution of drug-resistant strains of the virus.
Patrick Te Biology 1M03, 2011 The drug affected ordinary cells, but not the drug resistant mutant cells. Given an opportunity to proliferate, they begun to cause problems as the ordinary cells did. o Displays how individuals experience differential success and shows how evolution works through a change in allele frequency in a population, not through individuals. Galapagos Fiches (Peter and Rosemary Grant) o Researchers found that beak form and body size were heritable traits. During research, drought killed 84% of ground finches due to starvation. Natural selection resulting from this led to an increase in the average beak length. (Seed availability decreased, leaving large, hard fruits that were easier to break open for large, hard beaks). Another change of population characteristics occurred as a result of 7 months of rain. During this time, a reduction in average beak length occurred. (Shorter, pointier beaks are ideal for eating seeds, which were abundant). Selection vs. Evolutionary Change Natural selection acts on individuals, evolutionary change happens in the characteristics of the population. o Individuals do not change when selected, they simply survive and manage to reproduce with more frequency than other individuals. Acclimation occurs when an individuals phenotype (physical trait) is changed in response to environmental changes. The genotype (genetic trait) of an individual, however, remains fixed, so the changes made in acclimation are not passed on to offspring. Adaptation, on the other hand, has to do with allele frequencies in a population changing due to natural selection. As this has to do with genotypes, the traits are passed on to offspring. Clearing up some misconceptions: Evolution is NOT Goal-Directed or Progressive Evolution is not directed by a goal. It is simply a process that favors individuals that happen to be better adapted to an environment than others. Adaptations are not the result of a want or need. Evolution does not aim to produce better or more complex organisms. The idea of higher and lower organisms does not exist. Species are related by common ancestry and have evolved equally over time. Species may, however, become simpler or more complex depending on the environment. Animals Do Not Do Things for the Good of the Species Individuals with self-sacrificing alleles die and do not reproduce offspring. A possible exception to this fact is in kin-selection operation, but besides this, individuals do not sacrifice themselves for the good of the species. o Selfish alleles survive and produce offspring, therefore selfish alleles increase in frequency. Not all traits are adaptive. As adaptation is not a perfect process, the adaptations organisms have are constrained in a variety of ways. This includes genetic and historical constraints, and fitness tradeoff. Genetic Constraints o Selection is unable to optimize all aspects of a trait due to genetic constraints. o Genetic correlation occurs when selection favors alleles for one trait, but a correlated and suboptimal change occurs in an allele for another trait.
Patrick Te Biology 1M03, 2011 Lack of genetic variation can also constrain evolution because it works only on existing variation in a population. Fitness Trade-Off o A compromise between traits in terms of how they are adapted for an environment. o Selection acts on many traits at once, so an adaptation may be a compromise. Historical Constraints o When traits evolve from previously existing traits, adaptations are constrained by history. An adaptation made today was probably derived from a previous adaptation with a different application. The new application may not be the best solution for a given task. (ex. Bones in ear originally part of jaw) o Not all traits are adaptive. o
Chapter 25: Evolutionary Processes The Hardy-Weinberg principle acts as a model for generating predictions consistent with a null hypothesis when researchers wish to test whether one of 4 factors is affecting a given gene. o The four factors affect allele frequencies differently, with each having a different consequence. All four factors may not necessarily lead to adaptation. Natural selection produces adaptation in the way that it increases the frequency of certain alleles. Genetic drift produces stochastic (random) fluctuations in allele frequencies. Gene flow (migration) equalizes allele frequencies between populations. This results from individuals immigrating into or emigrating from a population. Mutation introduces new alleles. Continual introduction of new alleles eventually leads to a change in frequency. Biased or nonrandom mating, a 5th factor, can still affect evolutionary change. o Inbreeding, exemplified in selfing, changes genotype frequencies, but not allele frequencies. o Sexual selection leads to the evolution of traits that attract mates. It usually affects more strongly the traits of members in the gender that invests less in offspring and benefits from promiscuity, rather than the traits of the gender that will care for offspring and will thus benefit from choosy habits. The Hardy-Weinberg Principle (1908) Mathematical model to analyze consequences of mating among all individuals in a population. o Imagined that all of the gametes produced in a generation go into a gene pool before combining. o Resulting calculations predicted the genotypes of offspring produced, along with their frequency. Started with the simplest situation, a gene with alleles A1 and A2 o A1 frequency = p, A2 frequency = q, therefore in terms of a decimal percentage, p + q = 1 o Three varieties are possible in this case. A1A1, A1A2, A2A2. Review the Hardy-Weinberg principle as discussed in tutorial for the math stuff. :x The model predicts the following o The frequency of A1A1 will be equal to p2. o The frequency of A1A2 will be equal to 2pq.
Patrick Te Biology 1M03, 2011 The frequency of A2A2 will be equal to q2. P2 + 2pq + q2 = 1 When alleles are transmitted according to the rules of Mendelian inheritance (homozygous, heterozygous, etc), their frequencies do not change over time. Evolution occurs when other factors are taken into consideration. The Hardy-Weinberg Principle makes the assumption that none of the 5 factors leading to evolution is present in the population. The Hardy-Weinberg serves as a model for generating predictions consistent with a null hypothesis. This helps to determine whether a factor is acting ultimately on a particular gene or population. 1. Genotype frequencies that do not conform to Hardy-Weinberg proportions (equilibrium) indicate that a factor is affecting the population. To determine if the Hardy-Weinberg holds, you must 1. Estimate genotype frequencies by observation or testing. 2. Calculate observed allele frequencies from observed genotypes. 3. Use the observed allele frequencies to calculate the genotypes expected, assuming the population adheres to the Hardy-Weinberg. 4. Compare the observed and expected values. When the expected values do not match what is observed, we can assume that the one of the Hardy-Weinberg assumptions have been violated. o
Patterns of Natural Selection Natural selection occurs in a variety of patterns. Genetic variation refers to the number/relative frequency of alleles that are present in a particular population. o Maintenance of genetic variation is important because a lack of variations makes population less able to respond successfully to changes in the environment. Directional Selection o Pattern that increases the frequency of one allele by operating on a phenotype encoded at least partially by said allele. o Reduces genetic diversity over time. o As directional selection continues over time. Favored alleles approach a frequency of 1.0, while disadvantageous alleles approach a frequency of 0. The average distribution of a trait is changed. Alleles that reach a frequency of 1.0 (100%) are said to be fixed. Alleles no longer found in a population are said to be lost. Stabilizing Selection o Pattern found when individuals with intermediate traits reproduce more than others. o Stabilizing selection reduced genetic variation over time but does not affect the average value of a trait. Babies with birth weights in the middle of the average range are most likely to survive. Disruptive Selection o Contrasts to stabilizing selection; a pattern of disruptive selection is found when intermediate phenotypes are selected against, while extremes are favored. o Maintains / increases genetic variation but does not change the mean of a trait. o Can cause speciation if individuals with one extreme of a trait mate preferentially with individuals with the same trait.
Genetic Drift Genetic drift is any change in allele frequency in a population due to chance. Causes allele frequency to drift stochastically (element of randomness). Genetic drift is unbiased. These allele frequency changes are not adaptive. Over time, genetic drift can lead to stochastic loss or fixation of alleles. o Stochastic loss/fixation = genetic variation declines. More pronounced in small populations than large ones, but can still have an effect in large populations given time. Can be caused by any event or process that involves sampling error. Examples include o Founder Effect: Group immigrates to a new area and starts a population. If it is a small group, its allele frequencies likely differ from that of the source population. It is a sample of the old population that may not be reflective of the population. o Population Bottleneck: A sudden decrease in population size, usually caused by disease or natural disaster. Genetic Bottlenecks: Results from a population bottleneck, it defines a sudden reduction in the number of alleles in a population. Will occur when a disease or something of the type targets certain phenotypes. Gene Flow Gene flow, or migration, is the movement of alleles from one population to another and results when individuals leave one population to join and breed in another. Gene flow equalizes gene frequency between the source and recipient population. Gene flow may be unbiased with respect to fitness. However, allele movement between populations will always tend to reduce genetic difference between populations. Mutation Contrary to evolutionary mechanisms that reduce diversity, mutation restores it. Mutation is unbiased with respect to fitness. o This being said, the fact that mutations cause change in functioning genes mean that they often result in deleterious alleles that actually lower fitness. o It is only rarely that mutation produces a beneficial allele that increases in frequency within a given population due to natural selection. Mutation can be a significant force in bacteria due to their short generation times. Mutation tends to be slower in eukaryotes, however, rarely causing a significant change in genotype frequencies. Mutation, compared to other evolutionary mechanisms, is quite slow. Mutation likely introduces new alleles into every individual in every population in every generation, providing genetic variation that natural selection acts on in producing evolution. It is thus essential to the evolutionary process. Biased Mating In nature, mating may not always be unbiased with respect to a certain gene. Mechanisms of this include inbreeding and sexual selection. o Inbreeding: Mating between relatives. This increases the frequency of homozygotes in each generation. This does not cause allele frequencies to change in the population as a whole, though it does change genotype frequencies (increased homozygous).
Patrick Te Biology 1M03, 2011 Inbreeding depression is a decline in average fitness. This happens as a result of the increased homozygosity (and, as a result, decrease heterozygosity). Can increase the rate at which natural selection removes disadvantageous recessive alleles. Sexual Selection: Occurs when individuals in a population differ in their ability to attract mates. This violates Hardy-Weinberg assumptions because certain alleles are caused to increase or decrease in frequency, resulting in evolutionary change. Favors individuals with heritable traits that enhance ability to mate. Fundamental asymmetry of sex: females usually invest more time in offspring than males; therefore they should be choosier. They may choose on the bases of physical characteristics or resources. Male-male competition: Establishment of territory. Sexually selected traits often differ sharply between sexes. Sexual dimorphism defines any trait that differs between males and females of a single species.
Ch. 26: Speciation Key Concepts Speciation occurs when population of the same species become isolated due to a lack of gene flow, then diverge from each other due to any of the 5 factors, especially natural selection, genetic drift or mutation. Populations are recognized a distinct species if they are reproductively isolated from each other, have distinct morphological characteristics or form independent branches on a phylogenetic tree. o Genetic isolation can occur due to occupation of different geographic areas, different habitats, or if the species are unable to breed with one another. o Several outcomes are possible when diverged species come back into contact. Introduction Genetic divergence may eventually lead to speciation, or the creation of new species. Speciation tends to create two or more distinct species from a single ancestral group. Definition and Identification of Species A species is defined as an evolutionarily independent population(s). Identifying species can be difficult. Gene flow eliminates genetic differences among populations: evolutionary dependence starts with a lack of gene flow. o When gene flow between populations stop, the other 4 factors begin to act independently. o Allele frequencies and other characteristics of such population diverge, and can become a different species over time. Species are often identified in one of three ways The Biological Species Concept Considers populations to be evolutionarily independent if they are reproductively isolated from one another (no interbreeding). o No gene flow. Biologists categorize mechanisms that stop gene flow between two populations as prezygotic or postzygotic.
Patrick Te Biology 1M03, 2011 Prezygotic: Individuals of different species are prevented from mating. Temporal, habitat, behavior, gamete incompatibility or mechanical problems are all examples of this. o Postzygotic: Individuals from different populations can mate, but the hybrid offspring have low fitness and thus do not survive nor produce offspring. The hybrids are either not viable and die as embryos or are sterile for life. Invalid for fossils or creatures that reproduce asexually. Applies only to populations with a geographical overlap. The Morphospecies Concept Based on the idea that distinguishing features are most likely to arise if populations are independent and isolated from gene flow. o Biologists identify evolutionarily independent lineages by differences in size, shape or other such features. Wide application, but subjective. o Ex.) Are homo habilis and homo rudolfensis the same species or different species? The Phylogenetic Species Concept Based on the reconstruction of evolutionary history of a population. o An ancestral population + all its descendants constitute a monophyletic group or clade. A species is defined as the smallest monophyletic group on a tree containing populations. Each tip-set is a phylogenetic species. o
Applicable to any population, but currently available for only a tiny subset of populations on the tree of life. Biologists use all three species concepts. Isolation and Divergence in Allopatry Allopatric speciation involves physical isolation via dispersal or vicariance. o Populations living in different areas are said to be in allopatry. Biogeography is the study of how species and populations are distributed geographically, and can tell us how colonization and range-splitting events occur. Dispersal and Colonization Isolate Populations Colonization events often cause speciation due to the physical separation, reducing gene flow, and genetic drift (caused by the founder effect) causes the old and new populations to quickly diverge. o Natural selection may cause divergence if the newly colonized environment differs from the original habitat.
Patrick Te Biology 1M03, 2011 Vicariance Isolates Populations Glaciers serving as physical barriers during the last ice age is an example of a vicariance event. o Thought to be responsible for the origin of many modern species. Continental drift could have also separated species physically, though this would be a significantly larger-term process. o Ex.) Ratite birds (big birds) lived on Gondwana, which separated into South America, Australia, Africa etc. Ratite birds in each area are different, but probably came from a common ancestor. The physical isolation of populations produces genetic isolation, the first requirement of speciation. When combined with genetic divergence, speciation results. Isolation and Divergence in Sympatry Populations/species that live in the same geographic region live in sympatry. o Originally thought that speciation would not occur in sympatric populations because of gene flow. Under certain circumstances, sympatric speciation is in fact possible. Though not physically isolated, they may be isolated by preference or habitat. o Ex.) Soapberry bugs; some fed on one type of fruit, some the other. This caused a difference in the length of the average beak over time to match the fruit eaten. Disruptive selection (favored extremes) Natural selection overwhelms gene flow. Polyploidy and Speciation If populations become isolated, mutation alone is unlikely to cause an appreciable divergence. An exception is polyploidy (condition of having more than two sets of chromosomes), which, especially in plants, can cause speciation. o Ex.) Tetraploid individuals produce diploid gametes rather than haploid gametes. When combined with a haploid, a dysfunctional triploid results. Mutations that result in a doubling of chromosome number produce autopolyploid individuals. o All chromosomes come from the same species. Allopolyploid individuals are created when parents that belong to different species produce and offspring in which chromosome number doubles. Autopolyploidy Researchers found a population of tetraploid (4n) maidenhair ferns were the offspring of a parent that produced diploid gametes then self-fertilized. Polyploid populations such as these ferns are genetically isolated from wild-type populations because they cannot breed with diploids. o Genetic drift and natural selection cause a divergence between wild-type and polyploidy populations speciation is underway. Allopolyploidy May occur after the hybridization of two species. Many diploid plant species have closely related polyploidy species. o Supports the claim that speciation by polyploidy is important in plants. o Allopolyploidy speciation is driven by chromosome-level mutations and occurs in sympatry. When Isolated Populations Come into Contact If two populations have diverged and then come back into contact after divergence has affected when, where or how individuals mate, prezygotic isolation exists.
Patrick Te Biology 1M03, 2011 Mating between populations is rare, gene flow is minimal, the populations continue to diverge. If prezygotic isolation does not exist, populations may successfully interbreed. o Geneflow occurs, potentially erasing distinctions between the two populations. o Other outcomes could include reinforcement, development of hybrid zones and speciation by hybridization. Reinforcement If two populations of related species have diverged extensively and are distinct genetically, their hybrid offspring will most likely have a lower fitness and will not develop or reproduce normally. o Example of postzygotic isolation. o Indication of strong natural selection against interbreeding. Selection for traits that isolate populations reproductively is known as reinforcement. Sympatric species living in the same area tend to be seldom willing to mate with one another. Allopatric species living in different areas are conversely often willing to mate with one another. Hybrid Zones Sometimes, the hybrid offspring of related species possess traits that are intermediate between the two parental populations, and they are healthy and capable of breeding. Hybrid zones are geographic areas where interbreeding between two populations occurs. o Ex.) Warblers Hybridization often leads to extinction but sometimes leads to the origination of a new species. Hybridization Events There are a number of possible outcomes when secondary contact is made between two related populations o Fusion of the populations. o Reinforcement of divergence. o Founding of stable hybrid zones. o Extinction of one population. o Origination of a new species. Ch. 27: Phylogenies and the History of Life Key Concepts The Cambrian explosion was the rapid morphological and ecological diversification of animals that occurred during the Cambrian period. Adaptive radiations are a major pattern in the history of life. They are instances of rapid diversification associated with new ecological opportunities and new morphological innovations. Mass extinctions have occurred repeatedly throughout history. o Rapid elimination of most living species in an unbiased manner. Introduction Consider profound changes in the nature of life on Earth over large time periods. Two major analytical tools with which biologists reconstruct the history of life. Phylogenetic Trees A phylogeny is the evolutionary history of a group of organisms. o Phylogenetic trees show ancestor-descendant relationships among populations or species. o Ancestors and their descendants form a monophyletic group (clade/lineage). Branches represent groups through time. Adjacent branches are sister taxa. Tips are the endpoints of the tree, representing living groups or a groups extinction. o
Patrick Te Biology 1M03, 2011 Names at tips represent populations or higher taxa. Nodes occur where ancestral groups split into two or more decendant groups. o Polytomies are nodes where more than two decendant groups branch off. o In rooted phylogenies, the most ancient node of the tree is at the bottom. Location determined using an out-group, a taxonomic group that diverged before the rest of the taxa being studied. Estimating Phylogenies The analyzing of morphological and/or genetic characteristics allows one to infer phylogenetic relationships among species. There are two general strategies for using data to estimate trees: phenetic and cladistics. o The phenetic approach is based on a computing statistic that summarizes the overall similarity among populations. Computer program compares statistics for different populations, builds a tree that clusters most similar populations closer and more divergent species farther. o The cladistics approach focuses on synamorphies: the shared derived characteristics of species under study (trait in groups of organisms that do not exist in any others). When many synapmorphic traits are measured, traits unique to monophyletic groups are identified and the groups are placed on a tree in the most appropriate relationship to one another. Homology and Homoplasy Homology occurs when traits are similar due to shared ancestry. (ie. Genetics) Homoplasy occurs when traits are similar for reasons NOT related to common ancestry (ie. Similar living conditions natural selection has favored a similar solution to a problem posed by a similar way of life for two different creatures) Parsimony is a principle of logic that states that the most likely explanation or pattern is the one that implies the least amount of change. In the context of phylogenetic trees and changes in them, less is more. o If similar traits are found in distantly related relatives and is due to common ancestry, then it stands to reason that similar traits should be found in the species found on the tree of life between them. o Convergent evolution and other causes of homoplasy should be rare compared to cases of homology this would allow the tree to imply the fewest evolutionary changes. Fossils Physical traces left behind by past organisms, formed when an organism is buried in sediment before decomposition occurred. There are four types of fossils. o Intact fossils are formed when decomposition does not occur. o Compression fossils are formed when sediments accumulate on top of the fossil, pressing the organic material into a thin film. o Cast fossils result when decomposition happens after burial, with the space being filled by minerals. o Premineralized fossils are the result of very slow decay such that dissolved minerals gradually infiltrate the cells and harden into stone. The fossil record is the total collection of fossils that have been found throughout the world. o Provides the only direct evidence about what organisms that lived in the past looked like, where they lived, etc. Limitations of the Fossil Record
Patrick Te Biology 1M03, 2011 Habitat bias occurs because organisms that live in areas with the active deposit of sediments have a higher chance of becoming fossilized. Taxonomic bias is due to the fact that some organisms, such as those with bones, are more likely to decay slowly and leave fossil evidence behind. Temporal bias occurs because more recent fossils tend to be more common than more ancient ones due to the movement of tectonic plates, which may destroy deeper down / older fossils. Abundance bias occurs because organisms that are abundant, widespread and present on Earth for a long period will leave evidence of their existence far more often than species that are rare, local or short-lived. Lifes Time Line The Precambrian
The Paleozoic (Appearance of many animal groups)
The Mesozoic Age of Reptiles
The Cenozoic Age of Mammals
The Cambrian Explosion The first animals such as sponges, jellyfish and worms appear in the fossil record at around 565mya (end of the Proterozoic eon). o 50 million years later, these animals had diversified into almost all the major groups alive today. o Known as the Cambrian explosion period, arguably the time of the most evolutionary change in the history of life. An Overview of Cambrian Fossils Three major fossil assemblages. o Doushantuo, identified as tiny sponges, cyanobacteria, multicellular algae. 570-580mya. o Ediacaran, identified as sponges, jellyfish etc. (soft bodied). 544-565mya. o Burgess Shale, representing almost every major living animal group. 515-525mya. Gene Duplication and the Cambrian Explosion Many researchers predicted a strong association between the order in which animal lineages appeared during evolutionary history, the number of Hox genes present in each lineage, and each lineages morphological complexity and body size. o Hox genes help specify the location and shape of limbs, antennae, etc. o Logic behind new genes, new bodies hypothesis was that existing genes could have been duplicated before/during the Cambrian explosion to produce new body plans. Examining the cladygram, we can infer that o # of genes in Hox cluster expanded during course of evolution. o Genes appear to have been duplicated because the genes within the cluster are similarly structured and sequenced. o The entire cluster was duplicated several times in the lineage leading to vertebrates Both the duplication of Hox genes and changes in expression/function of existing genes were important. Adaptive Radiations Star phylogenies represent speciation events that were so rapid the order of branching could not be determined. Adaptive radiation is defined as rapid speciation in a single lineage followed by
Patrick Te Biology 1M03, 2011 divergence into many different adaptive forums. Can be triggered by ecological opportunity and morphological innovation. Ecological Opportunity as a Trigger One of the most consistent triggers of adaptive radiations is ecological opportunity, the availability of new types of resources. o Ex.) Lizards diversifying such that they are able to thrive in multiple environments that are available and will have less competition. Morphological Innovation as a Trigger Previously seen in the Cambrian explosion. Many of the other important diversification events in the history of life started with the evolution of one key trait that allowed for living in new areas, using new food sources, or moving in new ways. Mass Extinction The rapid extinction of a large number of lineages scattered throughout the tree of life. o >60% of species present are wiped out within 1 million years. Cause by catastrophic events. 5 recognized mass extinctions, the Big Five. Background extinction as opposed to mass extinction is a lower, average rate of extinction representing the normal loss of some species. Background Extinction Mass Extinction Typically occur when normal Result of extraordinary, sudden, environmental change, disease or temporary changes in the competition reduces populations to environment. zero. Mass extinctions function like genetic Thought to result from natural drift (random change in allele selection. frequencies). What Killed the Dinosaurs? The impact hypothesis for this phenomenon proposes that an asteroid struck Earth 65mya, causing the extinction of 60%-80% of existing multicellular species at the time. o Evidence in rock layers dated to 65mya, impact crater near Mexico. Selectivity Some lineages were better able to withstand environmental change due to the asteroid better than others. o Ex.) Dinosaurs and large marine reptiles died while mammals and amphibians survived. Recovery Very slow recovery after impact. Terrestrial ecosystems radically simplified, marine environment diversity remained low for 4-8 million years. o Mammals eventually diversified to fill niches left empty due to the mass extinction, and all major mammalian orders alive today appeared within 10-15million years. Chapter 50: An Introduction to Ecology Key Concepts Ecology investigates relationships between organisms and their environment. o Ecology provides the science necessary for the conservation of species and natural areas.
Patrick Te Biology 1M03, 2011 There are many branches of ecology, all meant to ask different questions. For example, consider, Why do birds sit on eggs? o Behavioral: What is the stimulus from the egg? o Physiological: What is responsible for the decision to sit? o Evolutionary: What happened in the evolution of birds that developed sitting? o Population: What are the consequences of sitting? One central goal of ecology is to understand the distribution and abundance of organisms. o Includes processes related to individual needs, social business, population and its impact, the food chain, etc.
Organismal Ecology Organismal ecologists look at what allows an individual organism to live successfully in a particular area. Population Ecology A population is a group of individuals of the same species that live in the same area at the same time. o Population ecologists investigate how the number of individuals in a population changes over time. Community Ecology A biological community consists of species that interact with one another within a given era. o Community ecologists study these interactions. o They also analyze how groups of species respond to natural disturbances, ex.) fire or flooding. Ecosystem Ecology An ecosystem consists of all the organisms living in a particular region. It also takes into account non-living/abiotic components. o Can be considered a concept that emphasizes aggregate properties by lumping species together. Ecosystem ecologists study interactions among these components and their physical aspects (ex. Energy flow) Ecological systems include aquatic and terrestrial systems. Physical Factors and Aquatic Environments The depth and movement of water are very important. Water depth dictates the availability of light. o Light in turn affects productivity; the total amount of carbon fixed by photosynthesis per unit area per year. Only certain wavelengths of light are available underwater. o Most light is blue-green. At 3m deep. The intensity of light declines with the depth of water. Pure seawater experiences a steeper negative correlation between depth and light intensity than pure freshwater. Types of Terrestrial Ecosystems Climate defines dominant vegetation types known as biomes. Climate itself is the prevailing, long term weather conditions found in an area. Weather is specific, short term atmospheric conditions of temperature, moisture, sunlight and wind.
Patrick Te Biology 1M03, 2011 o o Biomes are responses to climate and not weather. As such, biomes tend to occupy large areas of land. A vast variety of biomes are found around the world.
Productivity of Terrestrial Ecosystems On land, photosynthesis and plant growth are maximized when temperatures are warm and conditions are wet, for example in tropical rainforests. o Photosynthesis cannot occur effectively at low temperatures, such as in the Arctic tundra, nor under drought conditions, such as in the desert. Net primary productivity (NPP) is the total amount of carbon that is fixed per year, minus the amount of fixed carbon oxidized during cellular respiration. o Represents organic matter that is available as food for other organisms. o In terrestrial environments, NPP is often estimated by measuring aboveground biomass (the total mass of living plants, excluding roots). Ex.) Tropical wet forests feature high annual rainfall and warm conditions. Such conditions lead to very high biomass. Biomes To study biomes effectively, consider o Where biomes are relative to one another. o The typical climate of the biome. o The dominant type of vegetation. o Its special features, such as animal species. Subtropical Deserts 30N and 30S latitudes. o High average temperatures with moderate variation, low levels of precipitation. Scarcity of water. o Slow plant growth, low productivity. Plants are often widely spaced apart. Desert species adapt to the head and aridity by growing slowly or in spurts, especially after rainfall. Examples of prominent species include cacti. Temperate Grasslands Temperate regions. o Moderate temperatures, areas have long, warm summers and short, cold winters. The growing season is well defined. o Plants do not grow in winter. o Grasslands are dominated by grass which is relatively productive. Grasslands form when rain is less than required by forest. Temperate Forests Common biome in eastern North America, western Europe, east Asia, Chile, New Zealand. o Moderate average temperature and variation of temperature, areas have moderately high and relatively constant precipitation (low variance). Growing period stops momentarily during winter. Dominated by broad-leafed deciduous trees. Productivity is at lower levels than tropical rainforests but higher than deserts or grasslands. o Moderate species diversity. Boreal Forests The boreal forest or taiga forms just south of the Arctic Circle.
Patrick Te Biology 1M03, 2011 o Very cold winters, short, cool summers. Extreme temperature variation. Annual precipitation is low, but low temperatures limit evaporation, allowing the moisture of the area to remain high enough to support trees. Dominated by cold-tolerant conifers. o Low productivity, but aboveground biomass is high due to the long-lived, tall trees. Low species diversity. Arctic Tundra The arctic tundra in arctic regions where land is not covered in ice. o Cold, high temperature spread, low annual precipitation. The growing season lasts only 6-8 weeks, as temperatures are below freezing for most of the rest of the year. Most tundra soils are perennially frozen (permafrost layer). Dominated by small woody shrubs, lichens and herbaceous plants. o Low plant species diversity, low productivity, low aboveground biomass. Animal diversity is also low. The Role and Consequences of Climate Change Each type of aquatic environment and terrestrial biome has species adapted to the abiotic conditions present at a location. o Climate change profoundly affects these abiotic conditions. Global Patterns in Climate Change Tropics near the equator are warmer than areas farther than the equator due to the spread and intensity of light that hits them. Circulation cells exist at the equator. Warm air rises and cools, drops rain. As this happens, the cold air above gets pushed toward the poles. Conversely, warm air, lower to the ground, is pushed toward the equator (Hadley cell, also responsible for weather being so moist). Seasons are regular annual fluctuations in temperature or precipitation due to the earths tilt on its axis. Varying intensities of sunlight will hit a given area depending on this angle. o If Earth did not tilt on its axis, we would not have seasons! o As the Earth orbits the sun, different parts of it are hit at various angles. This is also why the seasons are inverted for north and south when we are receiving most of the sunlight and therefore heat (summer), Australia would be receiving the least (winter).
Mountains and Oceans Winds form oceans cool and drop precipitation on one side of a mountain range, but not the other side. o Mountain ranges cause the climates of either side of them to vary. o Rain shadow effect. Side with rain is forested and wet, other side is dry, desertlike. The oceans absorb heat from the atmosphere in summer and releases heat into the atmosphere during winter. o Oceans/other bodies of water moderate the temperatures of nearby land masses. o This is due to the high heat capacity of water when compared to air or the ground. Global Warming and Ecosystems Carbon dioxide, obviously, is the main contributing factor to global warming. Simulations, observations and experiments help us predict the effects of global warming. o Global warming is making climates more extreme. The amount of hurricanes, tornadoes, etc. may not change (frequency), but their intensity does. o Global warming is reducing species diversity. Studies have shown that less diversity exists at higher temperatures. Species Distribution Biogeography is the study of how organisms are distributed geographically. o The range (geographical distribution), of every species in Earth is limited. No organism can live everywhere. o Distribution can be understood through examining historical, biotic and physical conditions. The Role of History Dispersal is the movement of an individual from its place of birth to where it lives and breeds as an adult. o A lack of species in an area may indicate a physical barrier to its dispersal. Due to human activity in many cases. An exotic species introduced into an area may spread rapidly and eliminate native species. At this point, it becomes an invasive species. Biotic Factors The distribution of a species is usually limited by biotic factors including interaction with other organisms. o Ex.) Distribution of cattle is limited by tsetse flies that cause sleeping sickness, while the flies in turn depend on some types of vegetation and the presence of wild animals. Abiotic Factors Species distribution can also be limited by abiotic factors such as temperature and moisture. An area may be too cold or dry for an organism. o Often difficult to separate effects of biotic and abiotic factors on range of species.
Chapter 51: Behavior What is Behavior? All behaviors have two things in common. o They are actions of an organism. o They have some trigger or stimulus. o Ex.) Eating, swimming courting, building a nest, making tools, thinking or avoiding a test is behavior. Digesting is not. Key Ideas What are the ultimate and proximate causes of a behavior? That is, how does it happen at the genetic and physiological level, and how does it influence fitness? Within a single species, behavior ranges from highly stereotyped behavior to flexible, conditioned responses. Responses also vary between learned and unlearned. There are different types of learning. Methods of communication, orientation and navigation correlate closely with their habitat and the challenges that come with it. Altruistic behavior manifesting itself is usually due to helping relatives or helping individuals that will help them later. Ranges of Behavior A bird sitting on a test is highly stereotyped, an invariable response to eggs. Fashioning a tool in order to obtain something is a conditional, highly flexible response that changes through learning. o Learning is a change in behavior that results from a specific experience of an individual. o Behavioral traits vary on two axes. Innate behavior to behavior that originates through and is modified by learning. Highly stereotyped with little variation to highly flexible with a dependence on condition. Innate Behavior Fixed action patterns (FAPs) are stereotypical behavior patterns usually triggered by simple stimuli known as signal stimuli or releasers. o Example of innate behavior, that is, behavior that is inherited and varies little based on learning or the condition of the individual. o Most common when The impact on fitness is high and the response must be fast (reflex-like). Learning is not possible. o Ex.) Running in the face of a deadly threat, suckling in mammals, begging for food in bird hatchlings. o Ex2.) Bird song is innate. A bird reared with the song of a different bird will still for the most part follow the song normal for its species. Ex3.) How far do you know how to fly for food as a bird? It depends on the rewards. The maximization of feeding efficiency is known as optimal foraging. o You would go farther if doing so meant more food. If it didnt you would only go as far as the positive correlation lasted. Ex4.) The size advantage hypothesis states that if a dominant male in a group of bluehead wrass dies, the largest female in the group becomes male. o Energetically costly, but the female will have more offspring as a male fertilizing eggs than as a female laying eggs. o This is an example of genetics guiding behavior in a social context.
Learning Learning is of particular importance in species with large brains and lifestyles dominated by complex social interaction. o In species with large brains, FAPs and other types of inflexible, stereotyped behavior are rare. It is more common that each individual is capable of a wide range of behaviors. In classical conditioning, individuals are trained by experience to give a response to a stimulus that has nothing to do with the response. o Ex.) Pavlovs dogs being trained to salivate when hearing a bell ring, rather than salivating only when food is presented. Imprinting is a fast, irreversible kind of learning that only occurs during a short critical or sensitive period and lasts for life. o Ex.) Ducks and goslings adopt their mother as the first moving thing they see. o Ex2.) The ability of humans to learn a language shares common characteristics with imprinting. Can Animals Think? Several types of learning exist, and the ability to learn varies widely among species. Cognition (thinking) is the recognition and manipulation of facts about the world, along with the ability to form concepts and gain insights. o Crows can make tools and solve complex problems, suggesting that they are capable of cognition. Communication Communication is a process in which a signal (information-containing behavior) from one individual changes the behavior of another. o Communication is a social process. Communication is not simply the sending of a signal, but also the reception and acting upon a signal. o Ex.) The waggle dance of bees bees relay the location of food to the hive, which follows the bees direction. The type of signal tends to correlate with the environment of the creature. o Light does not travel very far through bodies of water, but sound does. Thus, many of the forms of communication found in the ocean are sound-based. o Light travels well in flat plains, so the signals of animals there are more visual. o Creatures that hunt at night may use scent to convey signals both effectively and quietly. Altruism Altruism is a behavior that has a fitness cost to the individual that exhibits it and a fitness benefit to the recipient. o While animals do not generally behave for the good of a species and individuals with self-sacrificing alleles tend to die and not produce offspring Kin selection is one example of a condition in which an animal may be inclined to behave altruistically. Reciprocal altruism refers to altruism among nonrelatives. Hamiltons Rule: B r > c o B defines the benefit to a recipient. o r is the coefficient of relatedness (fraction of alleles among the actor and beneficiary) r = in siblings, in half-siblings. o c represents the cost to the actor.
Patrick Te Biology 1M03, 2011 The tendency for altruistic behavior increases when
Ch. 52: Population Ecology Key Concepts Life tables are summaries of how likely it is that individuals of each age class in a population survive and reproduce. From these tables, the growth rate of a population can be calculated. A wide variety of patters of change in population size over time exist. Population ecology helps to predict the survival chances of endangered species and intervene if necessary. Demography The number of individuals present in a population depends on birth, death, immigration and emigration. We need to know the details of a populations makeup if we want to predict its future state. Such information includes o How many individuals of each age group are alive. o The probability of individuals in these age groups to survive until next year. o How many offspring are produced by females of different ages. It is also important to be aware of how many individuals, and of which ages, are immigrating or emigrating in a given generation. Life Tables As previously stated, life tables summarize the probability that an individual will survive and reproduce in any given year over its whole lifetime. Example: Year # alive Survivorship Fecundity Survivorship x fecundity = average # (born in same offspring produced per female time period) 0 1000 1 0.00 0.00 1 424 0.424 0.06 0.03 Survivorship This is a key component of a life table and is defined as the proportion of offspring that survive, on average, to a particular age. o There are various patterns of survivorship that can be shown as a survivorship curve. o Type 1: majority live until an old age, at which point survivorship drops drastically. o Type 2: survivorship steadily decreases as age increases. o Type 3: very low survivability from birth, very few live to see old age. Fecundity Another key component of the life table, it is defined as the number of female offspring produced by each female in the population. Age-specific fecundity is the average number of female offspring produced by a female in a given age class (specific age/age range). The Role of Life History
Patrick Te Biology 1M03, 2011 An organisms resources (time, energy) are limited. Organisms must choose whether to invest in some activities or other activities. o Organisms cannot have both high fecundity AND survival they must pick one. This is an example of a fitness trade off. o
Ex.) A female bird lays eggs: More eggs (high fecundity) means lower chance of survival for each individual. An organisms life history consists of traits determining how the organism allocates its resources to growth, reproduction and survival-related activities.
Population Growth A populations growth rate is the change in the number of individuals in a population (N) per unit of time (t). In the case that no immigration or emigration is taking place, the growth rate is equal to the population times the per-capita rate of increase. (growth rate = N x r) o The per capita rate of increase (r) is the difference between the birth and death rate. (r = b d) When births<deaths, r < 0, the population declines. When births>deaths, r > 0, the population grows. When birth rates per individual are maximized and death rates are minimized, r will be at a maximum. rmax is referred to as the intrinsic rate of increase. o Expressed as N / t = rmax x N o A given species has a characteristic rmax that does not change. However, the per-capita rate of increase of a population is usually much lower than the rmax value. o r values vary even between populations of the same species at the same time. They change over time for a given population as well. Exponential Growth Occurs when r does not change over time and is independent on the number of individuals in the population. o Growth is density independent when the increase in the size of a population does not affect r. o Growth is density dependent when the per-capita birth rate decreases and the percapita death rate increases as population density increases. (Survival and fecundity decline) o Speed of exponential growth dependent on r. Remember that N / t = r x N, therefore a large r value will yield a large population.
Logistic Growth Carrying capacity, K, refers to the maximum number of individuals in a population that can be supported in a particular habitat over a sustained period of time. It can change depending on conditions. If a population size of N is below the carrying capacity of K, the population should continue to grow, slowing down as the population size begins to increase and eventually hitting 0. o Proportional to (K-N)/K, and, N / t = rmax x N((K-N)/K) o This expression is known as the logistic growth equation and describes logistic population growth. The change in growth rate occurs as a function of the size of a populations size.
Population Dynamics Population dynamics is concerned with changes in populations through time and their causes. Natural populations show a range of patterns in addition to exponential and logistic growth. How do Metapopulations Change through Time? Metapopulations are populations of populations and the collectively make up a population that has been split between many small patches of habitat. Individual patches may follow logistic or exponential growth. Though some of these subpopulations can go extinct over time, migration can restore or reestablish these populations. o Overall, the whole population remains fairly stable. Why Do Some Populations Cycle? Some populations exhibit regular fluctuations in size. Cycles may have density dependent factors such as predation, disease or food shortages, which intensify as population density nears K and culminates in a population crash. Ex.) Hare-Lynx cycle o Hare density was lowest with lack of food and predators present. o Hare density was slightly higher without predators but still a lack of food. o Hare density was slightly higher in the presence of predators but an abundance of food. o Hare density far greater with a lack of predators and abundance of food. Therefore, population is limited by predation and food. CYCLE! Hare pop up => Lynx pop up => hare pop down => lynx pop down etc. How Does Age Structure Affect Population Growth? A populations age structure, that is, proportion of individuals at each possible age, has a dramatic effect on the populations growth over time. Ex.) The primrose; numbers and age are dependent on the amount of light they can get.
Patrick Te Biology 1M03, 2011 1. Populations dominated by juveniles should experience rapid growth. However, population size then declines with time due to the density-independent factor of shading due to trees. 2. The long-term trajectory of the population in an area depends primarily on the frequency/severity of circumstances that would create sunlit gaps (ex. Windstorms). This means that the dynamics of the population are governed by an abiotic, densityindependent factor. 3. In a large tract of forest, the population will be divided among several subpopulations each found in canopy gaps (metapopulation). Age Structure in Human Populations Human populations have been growing rapidly, and the growth rate was accelerating. This puts a strain on resources and will eventually have a very serious impact on human population. In developing countries, there is high fecundity but low survival. As developing countries develop, survival will increase and fecundity decrease, but the overall population will still increase due to the sheer number of younger adults that can produce offspring. Different fertility rates make a big difference mortality Is a minor factor in the growth of the massive human population. o When women produce exactly enough offspring to replace her and her offsprings father, zero population growth (ZPG) results. (r will = 0, births = deaths). o The future of the human population hinges on fertility rates. Chapter 53: Community Ecology Interactions among species have two main outcomes. o Affect the distribution and abundance of interacting species. o Such interactions are agents of natural selection (effect on evolution), and the outcomes of such interactions vary depending on many conditions. A biological community consists of interacting species that live in a defined area. Species Interactions The implication of interspecies interaction is that the population of one species may be linked to others that share its habitat. Biologists can analyze interactions among species by considering the effects on the fitness of involved individuals. o When a species benefits, it is a (+) interaction. o When a species is negatively impacted, it is a (-) interaction. o When a species is unaffected, it is a (0) interaction. Commensalism (+/0) is when one species indirectly affects the other for the better. Competition (-/-) Type Interaction Intraspecific competition refers to competition within species. Space, food, resources, etc. Interspecific competition refers to competition between species. Consumptive competition refers to when species consume the same resources. (ex. Water) Preemptive competition refers to when one species makes space unavailable to the other. (ex. Barnacles) Overgrowth competition refers to when one species grows above another. (ex. Tall trees)
Patrick Te Biology 1M03, 2011 Chemical competition refers to when one species produces toxins that negatively affect the other. (Ex. Salvia) Territorial competition refers to when a mobile species protects its feeding/breeding territory against others (ex. Different types of bears) Encounter competition refers to when two species interfere directly for access to specific resources. Niche Concept and Competition A niche can be thought of as the range of resources that a species is able to use, or the range of conditions that are tolerable. o Example: A species of bird will eat a seed of a certain size and avoid seeds that are much larger/much smaller than that seed. o Niche overlap occurs when interspecific competition for seeds of a certain size is present.
Niche Overlap and Fitness Competitive exclusion occurs when competition is asymmetric. When niches overlap completely, the species with the lower fitness is driven to extinction.
When niches do not overlap completely, however, the species with lower fitness may simply use resources that do not overlap with the stronger. (Weaker may evolve away from stronger). Fitness Trade-Offs
Patrick Te Biology 1M03, 2011 The use of a given resource is only one part of a niche. Individuals that are good competitors for certain resources are usually less able to deal with extremes in climate or sickness thus, there is a trade-off. o Ex.) Chthalamus grows slowly, but is protected well against predators and other dangers as it grows. o Ex)Semilanus grows fast, but is only really able to capture most of its space when life in general is easier.
Consumption and Parasitism(+/- ) Type Interaction Herbivory refers to the consumption of plants by herbivores. Parasitism refers to a parasite consuming relatively small amounts of tissue/nutrients from a host. The relationship is usually long term but not always fatal. Predation refers to predators killing and consuming another individual (prey). Standing or Constitutive Defenses These defenses are always present. Camouflage is blending into an environment. Schooling is travel in large numbers. Weaponry is the ability to combat potential predators. Mimicry can be employed by both dangerous and harmless species. o Mullerian mimics are genuinely dangerous. o Batesian mimics merely imitate actual dangerous species. Inducible Defenses These defenses are manifested only when needed. Ex.) Mussels in areas of high predation were more strongly attached to their base/substrate and had thicker shells than those in lower predation areas. Can Predators Reduce Prey Populations? o Example of wolf hunting resulting in tripled population of moose (pg1263) o Predator-prey cycle: when the population of the predator increases, the population of the prey tend to decrease as more are hunted. Why Dont Herbivores Eat Everything/Why is the World Green? This can be attributed to three main hypotheses. o The top-down control hypothesis, which states that predation/disease limits herbivore populations. o The poor-nutrition hypothesis, which states that plants are a poor food source in terms of nutrients provided (especially oxygen). This lack of nutrients limits the population. o The plant-defense hypothesis, which states that plants defend themselves (through thorns, poison, etc.) well enough to limit the populations of herbivores. Parasite Manipulation of Prey Parasites not only utilize the resources found in the body of a host, but may also manipulate their hosts in order to make transition to new hosts easier. Ex.) Flatworms.
Mutualism (+/+) Type Interaction Involves a wide variety of organisms and rewards. o Ex.) Flowers and bees. o Ex.) Fungi and plant roots (fungi receive sugars in exchange for N and P) Should not be confused with altruistic or self-sacrificing behavior; the benefits received by a species are simply a by-product of the other species pursuing its own interests.
SUMMARY OF INTERACTIONS Interaction Fitness Short-Term Impact: Distribution Type Effect and Abundance Competition -/Population of both sides reduced. In the case of asymmetric competition, the range of one species may be reduced. Consumption +/Impact to prey/host depends on and Parasitism prey density and how effectively they defend themselves.
Long Term Impact: Coevolution Niche differentiation (via natural selection) to reduce competition.
Mutualism
+/+
Commensalism
+/0
Population size and range of both species are dependent on one another. Population size and range of commensal may depends on size and distribution of host.
Strong selection applied to prey/host that favors effective defense. Strong selection applied to predator/parasite favors effective bypassing of defense. Strong selection on both species to maximize fitness benefits and minimize fitness costs of relationship. Strong selection on commensal to increase fitness benefits in relationship.
Community Structure While research on species interaction usually focuses on two species at a time, but biological communities may contain thousands of species the scope must be widened to fully understand the communitys structure. How Predictable are Communities? (Two Theories) Fredrick Clements: Communities develop by passing through a series of predictable stages. o Stages result from extensive interspecies interaction.
Patrick Te Biology 1M03, 2011 Continues changing and developing until reaching a final stable stage (climax community) Henry Gleason: Community found in a given area is neither stable nor predictable. o Whole process of developing is largely a matter of chance; more emphasis on abiotic factors and history of disturbances. Current and Past Species Distributions The past serves as evidence against entirely predictable communities! o After the glaciers retreated, groups of species did NOT change their geographical distributions in sync with one another, instead moving independently and at different speeds. o Fossilized pollen suggests that the composition of most plant communities has been dynamic and contingent on historical events rather than static or fixed by way of composition. Present experimentation has also partially disproven complete randomness. o 12 identical ponds with sterile water were compared after being left alone for a year. o Most species developed were the same across all ponds, but there were some species that were unique to some ponds. Therefore, we can say that both Clements and Gleason are both partially correct. While there is some order, Clements idea was too extreme. Gleasons view is more reflective of reality. Keystone Species The structure of a community can change dramatically if a single species of predator or herbivore is removed or added to a community. o Such a species is known as a keystone species they have a far greater impact on the surrounding species than their abundance would suggest. o
Community Dynamics The composition of communities changes over time. A disturbance is any event that removes some individuals or biomass from a given community. Usually, these disturbances follow predictable trends. A disturbance regime represents the impact of the disturbance and is determined by o Type of disturbance. o Frequency of disturbance. o Severity of disturbance. Succession: Development after Disturbance Succession refers to the recovery species make following the occurrence of a severe disturbance. o Primary succession occurs when a disturbance removes the soil and all organisms living in and above the soil. Volcanoes, glaciers and landslides do this.
Patrick Te Biology 1M03, 2011 Secondary succession occurs when a disturbance removes some or all organisms from an area but allows the soil to remain intact. Fires and logging do this. o The specific sequence of species that appears over time is called the successional pathway. Theoretical Considerations Biologists focus on three factors to predict the outcome of succession in a community. o The trait of the species evolved. o How the species interact. o Historical/environmental circumstances. Pioneering species (the first to arrive at a newly disturbed site) o Have good dispersal ability. o Are able to tolerate sever abiotic conditions. o Have high reproductive rates, but have little competitive ability. Once colonization is underway, the focus shifts from a capacity to deal with a harsh abiotic environment and more on how species interact with one another. o Facilitation occurs when early-arriving species make conditions more favorable for the arrival of later species. o Tolerance occurs when existing species do not affect the probability that subsequent species will become established. o Inhibition occurs when the presence of a species limits or inhibits the establishment of another. Species Richness in Ecological Communities Species richness is the number of species present in a given community. Species diversity is a weighted measure that incorporates a species relative abundance as well as its relative presence or absence. Predicting Species Richness: Island Biogeography Larger patches of habitat = more species than in smaller patches. o Islands in the ocean have a smaller number of species than do areas of the same spze on continents. The number of species present on an island is a product of two repeatedly occurring events: immigration and extinction. Species richness varies with three characteristics of an island: existing species, island size and remoteness, which together comprise the theory of island biogeophraphy. 1 2 3 o
In 1, we see that that as the number of species on an island increases, less species immigrate to the island and more become extinct. In 2, we see that larger islands have a larger rate of immigration and lower rate of extinction. The result of this is that larger islands tend to have more species. In 3, we see that islands near shore have a larger rate of immigration and lower rate of extinction. The result of this is that islands closer to other land masses tend to have more species.
Global Patterns in Species Richness Communities in the tropics have more species than communities in temperate or subarctic environments. o Negative correlation between latitude and species diversity. In attempting to explain this pattern, biologists have had to consider two principles. 1. Causal mechanism must be abiotic. 2. Species diversity of an area = speciation + immigration extinction emigration. There have been several hypotheses, none of which can fully define the observed pattern. Here are some of them. o The high-productivity hypothesis, which proposes that high productivity (total amount of photosynthesis), results in increased biomass which is consequently able to support a greater diversity of species. o The energy hypothesis, which proposes that high temperatures increase productivity, which increases diversity. It also increases the likelihood that organisms can tolerate the physical conditions of a region. o The time hypothesis, which proposes that tropical regions have had more time for speciation than other regions and thus will naturally have more diversity. o The intermediate disturbance hypothesis, which proposes that regions with a moderate type, frequency and severity of disturbance should have high species richness and diversity; the idea being that such regions will contain both pioneering species as well as species that will succeed longer-term. Chapter 54: Ecosystems Introduction An ecosystem consists of multiple communities of organisms that live in an area and their physical and chemical environments. Ecosystems have four components. 1. The abiotic environment 2. Primary producers 3. Consumers 4. Decomposers All of the above are linked by energy flow and nutrients. Land: Limited by warmth and moisture.
Patrick Te Biology 1M03, 2011 Water: Limited by nutrient availability. As energy flows from primary producers -> consumers -> decomposers, most of it is lost.
Energy Flow and Trophic Structure To analyze energy and nutrient flows, we focus on o The nature of the ecosystem compartments and where elements reside and on. o How quickly these elements move between reservoirs. The net primary productivity (NPP) is the energy that primary producers invest in producing new tissue. It represents the amount of energy available to consumers.
Global Patterns in Productivity NPP varies among ecosystems by region and biome. o Important to note that the most productive land ecosystems are located in the wet tropics and tends to decline as you head towards the poles, while marine productivity is highest along coastlines but otherwise does not seem to follow trends. What Limits Productivity? On land, a combination of temperature and the availability of water and sunlight (linked to features that define biomes!) limits productivity. In aquatic ecosystems, the availability of nutrients (phosphorus in freshwater, iron and nitrogen in oceans) limits productivity. How Does Energy Flow through an Ecosystem? NPP -> biomass, organic material that nonphotosynthetic organisms can consume. In all environments, the chemical energy in primary producers eventually moves to the next part of a food chain either primary consumers or primary decomposers. o Primary consumers are herbivores. o Primary decomposers include bacteria, fungi and archaea. Trophic levels refer to steps in energy transfer. Organisms that obtain their energy from the same type of source (ie. # steps away from primary producer) are on the same level.
Patrick Te Biology 1M03, 2011 A food chain connects the trophic levels in a particular ecosystem. o The grazing food chain is comprised of the network of herbivores and the organisms that consume them. o The decomposer food chain is made up of species that eat the dead remains of organisms. Food chains are usually embedded in more complex food webs. Food webs offer a more complete description of the trophic relationships among the organisms in an ecosystem, having many different and intertwining food chains. Why is Energy Lost at Each Trophic Level? All ecosystems exhibit a pyramid of production it is highest at the lowest level and declines as you go up trophic levels. 1. This pattern occurs because at each level, energy is lost and used for metabolism or other activity.
What Limits the Length of Food Chains? Food chains and food webs have only 2-7 trophic levels (avg of 4). Several hypotheses that try to explain this include 1. The energy-transfer hypothesis states that chain length is limited by productivity. Because productivity decreases significantly at each level, it would be very hard for individuals of populations at high levels to sustain themselves. 2. The stability hypothesis suggests that long food chains are easily disrupted by environmental factors. Disturbances in unstable environments may happen enough to result in food chains never reaching past a certain level before falling apart. 3. The environmental-complexity hypothesis states that food-chain length is a function of an ecosystems physical structure. 3-D environments like dense forests would accommodate more complex chains than, for example, a 2-D plain. Biogeochemical Cycles Biogeochemical cycles are paths that elements take as they move from abiotic systems through organisms and back again. There are three main things to consider. 1. The nature and size of reservoirs (where elements are stored for periods of time). 2. The speed at which the element moves between reservoirs and what influences this rate. 3. The interactions of the biogeochemical cycle with other cycles. Generalized Terrestrial Nutrient Cycle (Example) Deer eats leaves of a tree. Deers excretions / body upon death are broken down and consumed by decomposers. Nutrients yielded by the decomposers are taken up by trees. What Factors Control the Rate of Nutrient Cycling in Ecosystems?
Patrick Te Biology 1M03, 2011 Usually, it is decomposition of detritus that limits the rate at which nutrients cycle through an ecosystem. o Decomposition rate is influenced by abiotic conditions as well as the quality of the detritus used as a nutrient source. Deforestation also plays a role in nutrient loss. The presence of vegetation will limit the loss of nutrients, which get dissolved. Global Biochemical Cycles Nutrients leaving one ecosystem enter another. The movement of ions and molecules along ecosystems links local biogeochemical cycles into one massive system that stretches across the globe. The Global Water Cycle Water evaporates out of the ocean and precipitates back into it. (Evap > precip) Cycle completed by water that moves from land to the oceans via streams and groundwater. o Humans have deeply affected the water cycle, especially through groundwater, due to asphalt and concrete surfaces, which prevents replenishment of groundwater. Results in the lowering of the water table. The Global Carbon Cycle Largest reservoir of carbon is the ocean, followed by the atmosphere. Naturally, carbon moves in and out of organisms and between the various reservoirs. o The burning of fossil fuels has released an overabundance of carbon dioxide gas into the atmospheric pool of carbon dioxide from the inactive reservoir of the fossil fuel. o Contributes to global warming because carbon dioxide is a greenhouse gas (gas with heat trapping potential) that keeps heat on earth instead of allowing it to go back out into space. The Global Nitrogen Cycle The vast pool of molecular nitrogen N2 in the atmosphere is unavailable to plants. Nitrogen is only useful for the environment when it is reduced (fixed) into NH3. o Results from lightning-driven reactions in the atmosphere as well as from enzymecatalyzed reactions in bacteria that live in the soul and oceans. Humans artificially fix nitrogen in the form of fertilizers, burning fossil fuels and cultivating certain crops. o While nitrogen increases can be beneficial, nitrogen runoff (from fertilizers) can be disastrous for aquatic environments. It can also decrease biodiversity on land if species that respond more positively than other species become too dominant. Human Impacts on Ecosystems Human impact on ecosystems stems mainly from A rapid increase in human population. A rapid increase in human resource usage. In particular, residents of industrialized countries use a disproportionately large impact. Global Warming An increase in Earths surface temperature across the whole world. Current evidence has indicated that humans, mainly through an increase in greenhouse gases, are responsible. Examples of the implications of increased temperature o Changes in the geographic regions of many creatures (areas for warm-climate creatures expanding, cold-water species declining). Evolution is resulting from these changes.
Patrick Te Biology 1M03, 2011 o Coral reef bleaching. Productivity Changes Changes humans are inducing in cycles alter NPP. Generally, changes result in increasing NPP on land and decreasing NPP in oceans. Increased productivity can be bad decline in species richness, dead zones . o Dead zones result from decomposition of plankton taking up all the oxygen, leaving none for living organisms.

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Patrick Te Biology 1M03, 2011

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