4/3/13 Does life history affect molecular evolutionary rates?

Save this PDF as:
 WORD  PNG  TXT  JPG

Size: px
Start display at page:

Download "4/3/13 Does life history affect molecular evolutionary rates?"

Transcription

1 Does life history affect molecular evolutionary rates? By: Gerardo Antonio Cordero (Department of Ecology, Evolution and Organismal Biology, Iowa State University) & Fredric Janzen (Department of Ecology, Evolution and Organismal Biology, Iowa State University) 2013 Nature Education Citation: Cordero, G. A. & Janzen, F. (2013) Does life history affect molecular evolutionary rates? Nature Education Knowledge 4(4):1 The molecular evolutionary rate measures the frequency with which DNA or protein sequence mutations are fixed (i.e., shared by most individuals) in a population. On the other hand, the mutation rate refers to the amount of change in a DNA or protein sequence for a given unit of time. These two intrinsically related processes have been key to our basic understanding of molecular evolution. For example, the molecular clock theory (Zuckerkandl & Pauling 1962) predicts an increase in fixed amino acid mutations of protein sequences as a function of time. The molecular evolutionary rate is then expected to be constant, assuming that mutation rates are the same across the evolutionary lineages of a given protein (Figure 1). The neutral theory of molecular evolution (Kimura 1983) explains this pattern by proposing that most mutations do not have an effect on the fitness of an organism. Under this model, fixed DNA nucleotide mutations that do not result in changes to protein sequences are referred to as neutral or synonymous substitutions. Also, beneficial mutations are expected to be rare and deleterious ones quickly removed by natural selection. Although the molecular clock and neutral theories are foundational to modern molecular biology, they do not always explain why evolutionary rates vary among genes, proteins, and species. As a result, biologists have begun to consider how organismal-level traits such as life history could influence variation in molecular evolution across the tree of life. This is a central pursuit in evolutionary biology as it is relevant to key concepts such as speciation and the ability of an organism to adapt. 1/7

2 Figure 1: Under a molecular clock, fixed amino acid mutations of a protein sequence (vertical axis) increase as a function of time (hotizonal axis). Thus, the molecular evolutionary rate is expected to be constant (bold line). Each dot represents divergent evolutionary lineages of a given protein. Proteins in divergent evolutionary lineages that are not plotted on the line represent deviations from theory. Those to the left of the line are expected to have accelerated rates of evolution, while those to the right are expected to have decreased rates Nature Education Modified from concepts in Graur & Li All rights reserved. Molecular Evolution and Life history What determines variation in the mutation rate and ultimately the molecular evolutionary rate? The answer to this question is debatable for a number of reasons. Evidence at the cell and molecular levels clearly supports the finding that mutations often derive from DNA replication errors or mutagens in the environment. Organismal-level traits and population genetic processes, in turn, could influence the frequency of DNA replication errors and mutagen levels. Mutations are transmitted from generation to generation in the germ cell line of sexually reproducing organisms. Because DNA in germ cells replicates during meiosis (gamete differentiation), we expect short-lived species to have higher mutation rates. For example, mice 2/7

3 reproduce frequently and have short generation times. Their germ cell lines undergo more rounds of meiosis, thereby increasing the chances of DNA replication errors. Compared to longlived species, mice also are expected to have larger populations with more individuals that are available to reproduce that is, a larger effective population size. Consequently, synonymous (i.e., neutral) mutations are more likely to occur, leading to higher molecular evolutionary rates. Furthermore, the larger effective population size promotes selection against non-synonymous mutations that reduce fitness, thereby removing them from the population. Clearly, the nucleotide chains that comprise the nuclear and organellar genomes of eukaryotic cells do not evolve independently of organismal-level traits such as life history. Broadly speaking, life history refers to traits that function to regulate the life cycles of species (Roff 2002) (Figure 2). Examples of such traits include the timing of reproductive effort or its magnitude (fecundity). When is the best time to reproduce for an organism in a given habitat? How often does reproduction take place? These life-history traits are interrelated, making their potential effects on molecular evolutionary rates difficult to discern. However, biologists have proposed two models that establish a clear link between life history and molecular evolution: the generation time and metabolic rate hypotheses. The generation time hypothesis, illustrated in the mouse example, is related to the metabolic rate hypothesis because short-lived species with smaller body size usually have higher metabolic rates. Specifically, the metabolic rate hypothesis predicts that species with high metabolic rates have increased mutation rates due to mutagenic elements resulting from mitochondrial respiration (Galtier et al. 2009). Sometimes free oxygen radicals generated during respiration escape the mitochondrial electron transport chain that produces energy for the cell. These molecules cause damage to nearby mitochondrial DNA by oxidizing nucleotides. Figure 2: Organisms employ reproductive strategies that are adapted to their habitats. In this comparison, humans have a Type I survivorship curve in which a high percentage of offspring survive to adulthood and generation time is long. Frogs feature an opposite pattern (Type III) in which high numbers of offspring are produced but few survive. The small percentage that survives reaches adulthood quickly to produce 3/7

4 high offspring numbers. Birds display an intermediate strategy (Type II). Photo courtesy of Ray Husthwaite. To determine whether life-history traits such as generation time or metabolic rate affect molecular evolutionary rates, it is necessary to make comparisons across multiple evolutionary lineages. Molecular evolutionary rates and life-history differences among taxonomic groups are contrasted after accounting for shared common ancestry. The rate of molecular evolution is measured by using computer algorithms that estimate the number of substitutions (fixed mutations) per unit time in DNA sequences. This process is aided by using adequately dated fossils and well-supported hypotheses on the evolutionary relationship of species. We will briefly explore recent case studies that have adopted such approaches. Some of the most striking examples that illuminate the relationship between life history and molecular evolutionary rates are from plant and animal genomes. Life History and Molecular Evolution in Plants Differences in life-history strategies within taxonomic groups of flowering plants appear to influence molecular evolutionary rates (Figure 3). A comprehensive phylogenetic assessment of angiosperms found that rates of DNA sequence (chloroplast + mitochondrial + nuclear) evolution in long-lived (perennial) species are low compared to those of short-lived (annual) species (Smith & Donoghue 2008). This finding was consistent with the generation time hypothesis, and the results were subsequently validated by genome-wide assessments on model plant species. However, correlations were stronger in nuclear genomes as these featured higher molecular evolutionary rates than those of chloroplasts (Yue et al. 2010). Even when taking into account that some annual plant species are self-breeding, generation time still emerges as the strongest life-history correlate of molecular evolutionary rates (Muller & Albach 2010). Overall, the generation time hypothesis is the best-supported life-history model that explains molecular evolutionary rate variation in plants. There is little evidence to support the metabolic rate hypothesis in plants, although mitochondrial evolutionary rates are generally lower compared to those of animals. 4/7

5 Figure 3: Difference in the molecular evolutionary rate of annual (herbs) vs. perennial (trees) plants Nature Education (tree) Courtesy of Cordero. (flower) Photo via Wikimedia Commons. All rights reserved. Life History and Molecular Evolution in Animals Among mammals, mitochondrial evolutionary rates range from one substitution per 1-2 million years to one substitution per more than 100 million years a difference of 2 orders of magnitude (Nabholz et al. 2008A). Following the assumption that animal body size is correlated with metabolism, theoretical models strongly support the idea that molecular evolutionary rate variation is influenced by basal metabolic rate (Gillooly et al. 2005). In practice, support for this hypothesis is limited because few studies have addressed mutations in the germ cell line (Galtier et al. 2009). Germ cell line mutation rate was assessed in mutant strains of the nematode Caenorhabditis elegans with deficiencies in the mitochondrial electron transport chain. However, this experimental approach did not provide evidence for an increase in mitochondrial DNA mutation due to metabolic oxidative stress (Joyner-Matos et al. 2011). The relationship between life history and molecular evolutionary rates is more apparent in animal nuclear genomes. While a study on the effect of 14 life-history traits on molecular evolutionary rates of mitochondria in mammals did not provide strong support for the generation time hypothesis (Nabholz et al. 2008B), life history appears to influence molecular 5/7

6 evolution in the nuclear DNA of mammals (Bazin et al. 2006). Evidence suggests that their nuclear genomes evolve according to expectations from neutral theory. For example, the nuclear DNA of humans has a lower molecular evolutionary rate compared to primates that have shorter generation times. Specifically, in mammals, neutral evolutionary rates depend on generation time, while non-synonymous rates depend on population size (Nikolaev et al. 2007). Similar patterns have been demonstrated in invertebrate animals with the exception that generation time was correlated with non-synonymous rates (Thomas et al. 2010). These studies further support the generation time hypothesis, but we must keep in mind that it is not mutually exclusive from population size effects. Does life history drive molecular evolutionary rates? We have discussed evidence to support the idea that molecular evolutionary rates are driven by life history. By comparing differences among a wide variety of organisms, biologists can test the prediction that DNA nucleotide sequences do indeed evolve according to a rate that, at least partially, depends on organism-level traits. Generation time and metabolism, each to some degree or in combination, affect the mutation rates of some organisms and, thus, their molecular evolutionary rates. Even so, some relationships among generation time, metabolism, and molecular evolution depend on whether the organism is a plant or an animal and the location of the genome within the cell (i.e., nuclear vs. organellar). Also, differences in neutral vs. non-synonymous rates, when averaged together across long DNA sequences, could further complicate interpretations. Our understanding of variation in molecular evolutionary rates is likely to improve when more knowledge from molecular and organismal biology is made available. It is important to point out that the drivers of variation in molecular evolutionary rate even among genomic regions of a single species are not entirely understood. Work focusing on hypotheses concerned with DNA repair mechanisms, environmental effects, gene duplication, hypermutable nuclear DNA regions, population genetics, and sex-specific mutation bias may provide additional insights as to why rates of molecular evolution vary within and among species. An appreciation of these processes is necessary to clarify the origins of biological diversity and other evolutionary phenomena. References and Recommended Reading Bazin, E. et al. Population size does not influence mitochondrial genetic diversity in animals. Science 312, (2006). Galtier, N. et al. Mitochondrial whims: Metabolic rate, longevity and the rate of molecular evolution. Biology Letters 5, (2009). Gillooly, J. F. et al. The rate of DNA evolution: effects of body size and temperature on the molecular clock. Proceedings of the National Academy of Sciences (USA) 102, (2005). Graur, D., & Li, W.-H. Fundamentals of Molecular Evolution. Sunderland, MA: Sinauer Associates, Joyner-Matos, J. et al. No evidence of elevated germline mutation accumulation under oxidative stress in Caenorhabditis elegans. Genetics 189, (2011). Kimura, M. The Neutral Theory of Molecular Evolution. Cambridge, UK: Cambridge University Press, /7

7 Muller, K., & Albach, D.C. Evolutionary rates in Veronica L. (Plantaginaceae): Disentangling the influence of life history and breeding system. Journal of Molecular Evolution 70, (2010). Nabholz, B. et al. Strong variations of mitochondrial mutation rate across mammals-the longevity hypothesis. Molecular Biology and Evolution 25, (2008A). Nabholz, B. et al. Determination of mitochondrial genetic diversity in mammals. Genetics 178, (2008B). Nikolaev, S.I. et al. Life-history traits drive the evolutionary rates of mammalian coding and noncoding genomic elements. Proceedings of the National Academy of Sciences (USA) 104, (2007). Roff, D. Life History Evolution. Sunderland, MA: Sinauer Associates, Smith, S. A., & Donoghue, M. J. Rates of molecular evolution are linked to life history in flowering plants. Science 322, (2008). Thomas, J. A. et al. A generation time effect on the rate of molecular evolution in invertebrates. Molecular Biology and Evolution 27, (2010). Yue, J.-X. et al. Genome-wide investigation reveals high evolutionary rates in annual model plants. BMC Plant Biology 10, 242 (2010). Zuckerkandl, E. & Pauling, L. B. "Molecular disease, evolution, and genetic heterogeneity," in Horizons in Biochemistry, eds. M. Kasha & B. Pullman.(New York: Academic Press, 1962) /7

AP Biology Essential Knowledge Student Diagnostic

AP Biology Essential Knowledge Student Diagnostic AP Biology Essential Knowledge Student Diagnostic Background The Essential Knowledge statements provided in the AP Biology Curriculum Framework are scientific claims describing phenomenon occurring in

More information

Unit 1 - Fundamental Biology Skills and Knowledge

Unit 1 - Fundamental Biology Skills and Knowledge PREP TM AP* Biology Prep Course Syllabus Foundational Topics Review 10 units that cover fundamental biology topics typically covered in a general biology course. This content is perfect to use as a summer

More information

Principles of Evolution - Origin of Species

Principles of Evolution - Origin of Species Theories of Organic Evolution X Multiple Centers of Creation (de Buffon) developed the concept of "centers of creation throughout the world organisms had arisen, which other species had evolved from X

More information

Biology 1406 - Notes for exam 5 - Population genetics Ch 13, 14, 15

Biology 1406 - Notes for exam 5 - Population genetics Ch 13, 14, 15 Biology 1406 - Notes for exam 5 - Population genetics Ch 13, 14, 15 Species - group of individuals that are capable of interbreeding and producing fertile offspring; genetically similar 13.7, 14.2 Population

More information

CHAPTER 23 THE EVOLUTIONS OF POPULATIONS. Section B: Causes of Microevolution

CHAPTER 23 THE EVOLUTIONS OF POPULATIONS. Section B: Causes of Microevolution CHAPTER 23 THE EVOLUTIONS OF POPULATIONS Section B: Causes of Microevolution 1. Microevolution is generation-to-generation change in a population s allele frequencies 2. The two main causes of microevolution

More information

Nature of Genetic Material. Nature of Genetic Material

Nature of Genetic Material. Nature of Genetic Material Core Category Nature of Genetic Material Nature of Genetic Material Core Concepts in Genetics (in bold)/example Learning Objectives How is DNA organized? Describe the types of DNA regions that do not encode

More information

Chapter 21 Active Reading Guide The Evolution of Populations

Chapter 21 Active Reading Guide The Evolution of Populations Name: Roksana Korbi AP Biology Chapter 21 Active Reading Guide The Evolution of Populations This chapter begins with the idea that we focused on as we closed Chapter 19: Individuals do not evolve! Populations

More information

Chapter 16 Evolution of Populations. 16.1 Genes and Variation Biology Mr. Hines

Chapter 16 Evolution of Populations. 16.1 Genes and Variation Biology Mr. Hines Chapter 16 Evolution of Populations 16.1 Genes and Variation Biology Mr. Hines Figure 1-21 Levels of Organization Section 1-3 Levels of organization Biosphere Ecosystem The part of Earth that contains

More information

1 Mutation and Genetic Change

1 Mutation and Genetic Change CHAPTER 14 1 Mutation and Genetic Change SECTION Genes in Action KEY IDEAS As you read this section, keep these questions in mind: What is the origin of genetic differences among organisms? What kinds

More information

Ch. 13 How Populations Evolve Period. 4. Describe Lamarck s proposed theory of evolution, The Theory of Acquired Traits.

Ch. 13 How Populations Evolve Period. 4. Describe Lamarck s proposed theory of evolution, The Theory of Acquired Traits. Ch. 13 How Populations Evolve Name Period California State Standards covered by this chapter: Evolution 7. The frequency of an allele in a gene pool of a population depends on many factors and may be stable

More information

1. Why is mitosis alone insufficient for the life cycle of sexually reproducing eukaryotes?

1. Why is mitosis alone insufficient for the life cycle of sexually reproducing eukaryotes? Chapter 13: Meiosis and Sexual Life Cycles 1. Why is mitosis alone insufficient for the life cycle of sexually reproducing eukaryotes? 2. Define: gamete zygote meiosis homologous chromosomes diploid haploid

More information

Evolution and Darwin

Evolution and Darwin Evolution and Darwin Evolution The processes that have transformed life on earth from it s earliest forms to the vast diversity that characterizes it today. A change in the genes!!!!!!!! Old Theories of

More information

Bob Jesberg. Boston, MA April 3, 2014

Bob Jesberg. Boston, MA April 3, 2014 DNA, Replication and Transcription Bob Jesberg NSTA Conference Boston, MA April 3, 2014 1 Workshop Agenda Looking at DNA and Forensics The DNA, Replication i and Transcription i Set DNA Ladder The Double

More information

How Populations Evolve

How Populations Evolve How Populations Evolve Darwin and the Origin of the Species Charles Darwin published On the Origin of Species by Means of Natural Selection, November 24, 1859. Darwin presented two main concepts: Life

More information

MCAS Biology. Review Packet

MCAS Biology. Review Packet MCAS Biology Review Packet 1 Name Class Date 1. Define organic. THE CHEMISTRY OF LIFE 2. All living things are made up of 6 essential elements: SPONCH. Name the six elements of life. S N P C O H 3. Elements

More information

Ch.16-17 Review. Name: Class: Date: Multiple Choice Identify the choice that best completes the statement or answers the question.

Ch.16-17 Review. Name: Class: Date: Multiple Choice Identify the choice that best completes the statement or answers the question. Name: Class: Date: Ch.16-17 Review Multiple Choice Identify the choice that best completes the statement or answers the question. 1. Which of the following statements describe what all members of a population

More information

PRINCIPLES OF POPULATION GENETICS

PRINCIPLES OF POPULATION GENETICS PRINCIPLES OF POPULATION GENETICS FOURTH EDITION Daniel L. Hartl Harvard University Andrew G. Clark Cornell University UniversitSts- und Landesbibliothek Darmstadt Bibliothek Biologie Sinauer Associates,

More information

Quiz #4 Ch. 4 Modern Evolutionary Theory

Quiz #4 Ch. 4 Modern Evolutionary Theory Physical Anthropology Summer 2014 Dr. Leanna Wolfe Quiz #4 Ch. 4 Modern Evolutionary Theory 1. T/F Evolution by natural selection works directly on individuals, transforming populations. 2. T/F A genotypic

More information

Mechanisms of Evolution

Mechanisms of Evolution page 2 page 3 Teacher's Notes Mechanisms of Evolution Grades: 11-12 Duration: 28 mins Summary of Program Evolution is the gradual change that can be seen in a population s genetic composition, from one

More information

Worksheet - COMPARATIVE MAPPING 1

Worksheet - COMPARATIVE MAPPING 1 Worksheet - COMPARATIVE MAPPING 1 The arrangement of genes and other DNA markers is compared between species in Comparative genome mapping. As early as 1915, the geneticist J.B.S Haldane reported that

More information

Essentials of Human Anatomy & Physiology 11 th Edition, 2015 Marieb

Essentials of Human Anatomy & Physiology 11 th Edition, 2015 Marieb A Correlation of Essentials of Human Anatomy Marieb To the Next Generation Science Standards Life A Correlation of, HS-LS1 From Molecules to Organisms: Structures and Processes HS-LS1-1. Construct an explanation

More information

Practice Questions 1: Evolution

Practice Questions 1: Evolution Practice Questions 1: Evolution 1. Which concept is best illustrated in the flowchart below? A. natural selection B. genetic manipulation C. dynamic equilibrium D. material cycles 2. The diagram below

More information

AP Biology 2015 Free-Response Questions

AP Biology 2015 Free-Response Questions AP Biology 2015 Free-Response Questions College Board, Advanced Placement Program, AP, AP Central, and the acorn logo are registered trademarks of the College Board. AP Central is the official online home

More information

What two Assumptions did Darwin have to arrive at BEFORE he could form his theories of evolution?

What two Assumptions did Darwin have to arrive at BEFORE he could form his theories of evolution? Influences on Darwin s Thinking: What ideas did each of the listed names below contribute to Darwin s thinking about evolution? (very brief) Georges Buffon: Jean Baptiste Lamarck: Charles Lyell: Thomas

More information

Summary. 16 1 Genes and Variation. 16 2 Evolution as Genetic Change. Name Class Date

Summary. 16 1 Genes and Variation. 16 2 Evolution as Genetic Change. Name Class Date Chapter 16 Summary Evolution of Populations 16 1 Genes and Variation Darwin s original ideas can now be understood in genetic terms. Beginning with variation, we now know that traits are controlled by

More information

AP BIOLOGY 2010 SCORING GUIDELINES (Form B)

AP BIOLOGY 2010 SCORING GUIDELINES (Form B) AP BIOLOGY 2010 SCORING GUIDELINES (Form B) Question 2 Certain human genetic conditions, such as sickle cell anemia, result from single base-pair mutations in DNA. (a) Explain how a single base-pair mutation

More information

LAB 11 Natural Selection (version 2)

LAB 11 Natural Selection (version 2) LAB 11 Natural Selection (version 2) Overview In this laboratory you will demonstrate the process of evolution by natural selection by carrying out a predator/prey simulation. Through this exercise you

More information

LECTURE 6 Gene Mutation (Chapter 16.1-16.2)

LECTURE 6 Gene Mutation (Chapter 16.1-16.2) LECTURE 6 Gene Mutation (Chapter 16.1-16.2) 1 Mutation: A permanent change in the genetic material that can be passed from parent to offspring. Mutant (genotype): An organism whose DNA differs from the

More information

Regents Biology REGENTS REVIEW: PROTEIN SYNTHESIS

Regents Biology REGENTS REVIEW: PROTEIN SYNTHESIS Period Date REGENTS REVIEW: PROTEIN SYNTHESIS 1. The diagram at the right represents a portion of a type of organic molecule present in the cells of organisms. What will most likely happen if there is

More information

Name Class Date. KEY CONCEPT Mutations are changes in DNA that may or may not affect phenotype. frameshift mutation

Name Class Date. KEY CONCEPT Mutations are changes in DNA that may or may not affect phenotype. frameshift mutation Unit 7 Study Guide Section 8.7: Mutations KEY CONCEPT Mutations are changes in DNA that may or may not affect phenotype. VOCABULARY mutation point mutation frameshift mutation mutagen MAIN IDEA: Some mutations

More information

Evolution (18%) 11 Items Sample Test Prep Questions

Evolution (18%) 11 Items Sample Test Prep Questions Evolution (18%) 11 Items Sample Test Prep Questions Grade 7 (Evolution) 3.a Students know both genetic variation and environmental factors are causes of evolution and diversity of organisms. (pg. 109 Science

More information

9.1: Mechanisms of Evolution and Their Effect on Populations pg. 350-359

9.1: Mechanisms of Evolution and Their Effect on Populations pg. 350-359 9.1: Mechanisms of Evolution and Their Effect on Populations pg. 350-359 Key Terms: gene flow, non-random mating, genetic drift, founder effect, bottleneck effect, stabilizing selection, directional selection

More information

2. is a process of nuclear division in which the number of chromosomes in certain cells is halved during gamete formation.

2. is a process of nuclear division in which the number of chromosomes in certain cells is halved during gamete formation. Meiosis 1. P. J. van Beneden proposed that an egg and a sperm, each containing half the complement of chromosomes found in somatic cells, fuse to produce a single cell called a. 2. is a process of nuclear

More information

Keystone Biology Exam Information: Module A: Cell and Cell Processes

Keystone Biology Exam Information: Module A: Cell and Cell Processes Keystone Biology Exam Information: Module A: Cell and Cell Processes Basic Biological Principles- Day 1 Describe the characteristics of life shared by prokaryotic and eukaryotic organisms. Compare cellular

More information

Understanding by Design. Title: BIOLOGY/LAB. Established Goal(s) / Content Standard(s): Essential Question(s) Understanding(s):

Understanding by Design. Title: BIOLOGY/LAB. Established Goal(s) / Content Standard(s): Essential Question(s) Understanding(s): Understanding by Design Title: BIOLOGY/LAB Standard: EVOLUTION and BIODIVERSITY Grade(s):9/10/11/12 Established Goal(s) / Content Standard(s): 5. Evolution and Biodiversity Central Concepts: Evolution

More information

Respiration occurs in the mitochondria in cells.

Respiration occurs in the mitochondria in cells. B3 Question Which process occurs in the mitochondria in cells? Why do the liver and muscle cells have large number of mitochondria? What is the function of the ribosomes? Answer Respiration occurs in the

More information

Evolution Part 1. Unit 10 Miss Wheeler

Evolution Part 1. Unit 10 Miss Wheeler Evolution Part 1 Unit 10 Miss Wheeler Evolution Evolution- The process by which organisms have changed (and will continue changing) over time Charles Darwin- Father of Evolution Traveled for 5 years on

More information

Worksheet: The theory of natural selection

Worksheet: The theory of natural selection Worksheet: The theory of natural selection Senior Phase Grade 7-9 Learning area: Natural Science Strand: Life and living Theme: Biodiversity, change and continuity Specific Aim 1: Acquiring knowledge of

More information

Let s get started. So, what is science?

Let s get started. So, what is science? Let s get started So, what is science? Well Science Science is the observation of phenomena and the theoretical explanation of it. Simply, it is the state of knowing. Biology Biology is the study of life.

More information

AS Biology Unit 2 Key Terms and Definitions. Make sure you use these terms when answering exam questions!

AS Biology Unit 2 Key Terms and Definitions. Make sure you use these terms when answering exam questions! AS Biology Unit 2 Key Terms and Definitions Make sure you use these terms when answering exam questions! Chapter 7 Variation 7.1 Random Sampling Sampling a population to eliminate bias e.g. grid square

More information

Science 10-Biology Activity 14 Worksheet on Sexual Reproduction

Science 10-Biology Activity 14 Worksheet on Sexual Reproduction Science 10-Biology Activity 14 Worksheet on Sexual Reproduction 10 Name Due Date Show Me NOTE: This worksheet is based on material from pages 367-372 in Science Probe. 1. Sexual reproduction requires parents,

More information

Problem Set 5 BILD10 / Winter 2014 Chapters 8, 10-12

Problem Set 5 BILD10 / Winter 2014 Chapters 8, 10-12 Chapter 8: Evolution and Natural Selection 1) A population is: a) a group of species that shares the same habitat. b) a group of individuals of the same species that lives in the same general location

More information

An Overview of Cells and Cell Research

An Overview of Cells and Cell Research An Overview of Cells and Cell Research 1 An Overview of Cells and Cell Research Chapter Outline Model Species and Cell types Cell components Tools of Cell Biology Model Species E. Coli: simplest organism

More information

C1. A gene pool is all of the genes present in a particular population. Each type of gene within a gene pool may exist in one or more alleles.

C1. A gene pool is all of the genes present in a particular population. Each type of gene within a gene pool may exist in one or more alleles. C1. A gene pool is all of the genes present in a particular population. Each type of gene within a gene pool may exist in one or more alleles. The prevalence of an allele within the gene pool is described

More information

Unit I: Introduction To Scientific Processes

Unit I: Introduction To Scientific Processes Unit I: Introduction To Scientific Processes This unit is an introduction to the scientific process. This unit consists of a laboratory exercise where students go through the QPOE2 process step by step

More information

Animal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri

Animal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri Animal Models of Human Behavioral and Social Processes: What is a Good Animal Model? Dario Maestripieri Criteria for assessing the validity of animal models of human behavioral research Face validity:

More information

AP Biology Unit I: Ecological Interactions

AP Biology Unit I: Ecological Interactions AP Biology Unit I: Ecological Interactions Essential knowledge 1.C.1: Speciation and extinction have occurred throughout the Earth s history. Species extinction rates are rapid at times of ecological stress.

More information

Chapter 16 How Populations Evolve

Chapter 16 How Populations Evolve Title Chapter 16 How Populations Evolve Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Population Genetics A population is all of the members of a single species

More information

Name Date. Meiosis Worksheet

Name Date. Meiosis Worksheet Name Date Meiosis Worksheet Identifying Processes On the lines provided, order the different stages of meiosis I THROUGH meiosis II, including interphase in the proper sequence. 1. homologous chromosome

More information

Name: Class: Date: Multiple Choice Identify the choice that best completes the statement or answers the question.

Name: Class: Date: Multiple Choice Identify the choice that best completes the statement or answers the question. Name: Class: Date: Chapter 17 Practice Multiple Choice Identify the choice that best completes the statement or answers the question. 1. The correct order for the levels of Linnaeus's classification system,

More information

A Correlation of Pearson Miller & Levine Biology 2014 To the Utah Core State Standards for Biology Grades 9-12

A Correlation of Pearson Miller & Levine Biology 2014 To the Utah Core State Standards for Biology Grades 9-12 A Correlation of Pearson To the Utah Core State Standards Resource Title: Publisher: Pearson Education publishing as Prentice Hall ISBN (10 or 13 digit unique identifier is required): SE: 9780133242003

More information

Theories of Evolution: A Brief History (take notes from classmates presentations)

Theories of Evolution: A Brief History (take notes from classmates presentations) Packet Theories of : A Brief History (take notes from classmates presentations) Carl Linnaeus (1707-1778) William Paley (1743-1805) Georges Cuvier (1769-1832) Thomas Malthus (1766-1834) Jean Baptiste Lamarck

More information

Name Class Date. binomial nomenclature. MAIN IDEA: Linnaeus developed the scientific naming system still used today.

Name Class Date. binomial nomenclature. MAIN IDEA: Linnaeus developed the scientific naming system still used today. Section 1: The Linnaean System of Classification 17.1 Reading Guide KEY CONCEPT Organisms can be classified based on physical similarities. VOCABULARY taxonomy taxon binomial nomenclature genus MAIN IDEA:

More information

AP Biology Syllabus 2012-2013

AP Biology Syllabus 2012-2013 n AP Biology, an emphasis is on students making connections between the big ideas within the AP Biology Curriculum Framework. he two main goals of AP Biology are to help students develop a conceptual framework

More information

A. Homologous chromosomes divide in Meiosis l and sister. B. Homologous chromosomes divide in Meiosis ll and sister

A. Homologous chromosomes divide in Meiosis l and sister. B. Homologous chromosomes divide in Meiosis ll and sister SC.912.L.16.17 1) Somatic cells undergo mitosis whereas gamete cells undergo meiosis. Mitosis takes place throughout the lifetime of an organism. What is the biggest difference between these processes?

More information

Biology 160 Lab Module 10 Meiosis Activity & Mendelian Genetics

Biology 160 Lab Module 10 Meiosis Activity & Mendelian Genetics Name Biology 160 Lab Module 10 Meiosis Activity & Mendelian Genetics Introduction During your lifetime you have grown from a single celled zygote into an organism made up of trillions of cells. The vast

More information

Microevolution is a change in population s gene pool [1]

Microevolution is a change in population s gene pool [1] GUIDED READING - Ch. 14 - Section 4 NAME: Please print out these pages and HANDWRITE the answers directly on the printouts. Typed work or answers on separate sheets of paper will not be accepted. Importantly,

More information

Replication Study Guide

Replication Study Guide Replication Study Guide This study guide is a written version of the material you have seen presented in the replication unit. Self-reproduction is a function of life that human-engineered systems have

More information

2.3: Eukaryotic Evolution and Diversity pg. 67. For about 1.5 billion years Prokaryotes were on the only living organism on Earth.

2.3: Eukaryotic Evolution and Diversity pg. 67. For about 1.5 billion years Prokaryotes were on the only living organism on Earth. 2.3: Eukaryotic Evolution and Diversity pg. 67 For about 1.5 billion years Prokaryotes were on the only living organism on Earth. 3.5 to 2 billion years ago Prokaryotes thrive in many different environments.

More information

A CONTENT STANDARD IS NOT MET UNLESS APPLICABLE CHARACTERISTICS OF SCIENCE ARE ALSO ADDRESSED AT THE SAME TIME.

A CONTENT STANDARD IS NOT MET UNLESS APPLICABLE CHARACTERISTICS OF SCIENCE ARE ALSO ADDRESSED AT THE SAME TIME. Biology Curriculum The Georgia Performance Standards are designed to provide students with the knowledge and skills for proficiency in science. The Project 2061 s Benchmarks for Science Literacy is used

More information

Biological Sciences Initiative. Human Genome

Biological Sciences Initiative. Human Genome Biological Sciences Initiative HHMI Human Genome Introduction In 2000, researchers from around the world published a draft sequence of the entire genome. 20 labs from 6 countries worked on the sequence.

More information

Evolution, Natural Selection, and Adaptation

Evolution, Natural Selection, and Adaptation Evolution, Natural Selection, and Adaptation Nothing in biology makes sense except in the light of evolution. (Theodosius Dobzhansky) Charles Darwin (1809-1882) Voyage of HMS Beagle (1831-1836) Thinking

More information

Okami Study Guide: Chapter 3 1

Okami Study Guide: Chapter 3 1 Okami Study Guide: Chapter 3 1 Chapter in Review 1. Heredity is the tendency of offspring to resemble their parents in various ways. Genes are units of heredity. They are functional strands of DNA grouped

More information

Localised Sex, Contingency and Mutator Genes. Bacterial Genetics as a Metaphor for Computing Systems

Localised Sex, Contingency and Mutator Genes. Bacterial Genetics as a Metaphor for Computing Systems Localised Sex, Contingency and Mutator Genes Bacterial Genetics as a Metaphor for Computing Systems Outline Living Systems as metaphors Evolutionary mechanisms Mutation Sex and Localized sex Contingent

More information

LAB 21 Using Bioinformatics to Investigate Evolutionary Relationships; Have a BLAST!

LAB 21 Using Bioinformatics to Investigate Evolutionary Relationships; Have a BLAST! LAB 21 Using Bioinformatics to Investigate Evolutionary Relationships; Have a BLAST! Introduction: Between 1990-2003, scientists working on an international research project known as the Human Genome Project,

More information

Student name ID # 2. (4 pts) What is the terminal electron acceptor in respiration? In photosynthesis? O2, NADP+

Student name ID # 2. (4 pts) What is the terminal electron acceptor in respiration? In photosynthesis? O2, NADP+ 1. Membrane transport. A. (4 pts) What ion couples primary and secondary active transport in animal cells? What ion serves the same function in plant cells? Na+, H+ 2. (4 pts) What is the terminal electron

More information

Meiosis and Sexual Life Cycles

Meiosis and Sexual Life Cycles Meiosis and Sexual Life Cycles Chapter 13 1 Ojectives Distinguish between the following terms: somatic cell and gamete; autosome and sex chromosomes; haploid and diploid. List the phases of meiosis I and

More information

SCORING KEY AND RATING GUIDE

SCORING KEY AND RATING GUIDE FOR TEACHERS ONLY The University of the State of New York Le REGENTS HIGH SCHOOL EXAMINATION LIVING ENVIRONMENT Tuesday, August 13, 2002 12:30 to 3:30 p.m., only SCORING KEY AND RATING GUIDE Directions

More information

This is JEOPARDY!!!! 7 TH Grade Life Science

This is JEOPARDY!!!! 7 TH Grade Life Science This is JEOPARDY!!!! 7 TH Grade Life Science Blind Me with Science Questions about the Nature of Science They Might Be Giant Cells Questions about Cells, Molecules, and Atoms Head, Shoulders, Knees and

More information

AP BIOLOGY SUMMER WORK 2016

AP BIOLOGY SUMMER WORK 2016 AP BIOLOGY SUMMER WORK 2016 Welcome to AP Biology! AP Biology is a rigorous course designed to be equivalent to a first year biology course at a university. This summer work was designed to get you started

More information

12.1 The Role of DNA in Heredity

12.1 The Role of DNA in Heredity 12.1 The Role of DNA in Heredity Only in the last 50 years have scientists understood the role of DNA in heredity. That understanding began with the discovery of DNA s structure. In 1952, Rosalind Franklin

More information

Heredity - Patterns of Inheritance

Heredity - Patterns of Inheritance Heredity - Patterns of Inheritance Genes and Alleles A. Genes 1. A sequence of nucleotides that codes for a special functional product a. Transfer RNA b. Enzyme c. Structural protein d. Pigments 2. Genes

More information

Theory of Evolution. A. the beginning of life B. the evolution of eukaryotes C. the evolution of archaebacteria D. the beginning of terrestrial life

Theory of Evolution. A. the beginning of life B. the evolution of eukaryotes C. the evolution of archaebacteria D. the beginning of terrestrial life Theory of Evolution 1. In 1966, American biologist Lynn Margulis proposed the theory of endosymbiosis, or the idea that mitochondria are the descendents of symbiotic, aerobic eubacteria. What does the

More information

Exam 1. CSS/Hort 430. 2008 All questions worth 2 points

Exam 1. CSS/Hort 430. 2008 All questions worth 2 points Exam 1. CSS/Hort 430. 2008 All questions worth 2 points 1. A general definition of plants is they are eukaryotic, multi-cellular organisms and are usually photosynthetic. In this definition, eukaryotic

More information

Chapter. Extranuclear Inheritance. Lecture Presentation by Dr. Cindy Malone, California State University Northridge Pearson Education, Inc.

Chapter. Extranuclear Inheritance. Lecture Presentation by Dr. Cindy Malone, California State University Northridge Pearson Education, Inc. Chapter 9 Extranuclear Inheritance Lecture Presentation by Dr. Cindy Malone, California State University Northridge Chapter 9: Introduction Inheritance Observations revealed inheritance patterns fail to

More information

Microevolution: The mechanism of evolution

Microevolution: The mechanism of evolution Microevolution: The mechanism of evolution What is it that evolves? Not individual organisms Populations are the smallest units that evolve Population: members of a species (interbreeding individuals and

More information

Lab 10 Mitosis. Background. Mitosis. Prokaryotic fission. Prophase During prophase, the chromatin. Eukaryotic cell division

Lab 10 Mitosis. Background. Mitosis. Prokaryotic fission. Prophase During prophase, the chromatin. Eukaryotic cell division Lab 10 Mitosis Background Reproduction means producing a new organism from an existing organism. The new offspring must receive hereditary information and enough cytoplasmic material to maintain its own

More information

Mutations & DNA Technology Worksheet

Mutations & DNA Technology Worksheet Mutations & DNA Technology Worksheet Name Section A: Mutations Mutations are changes in DNA. Somatic mutations occur in non-reproductive cells and won't be passed onto offspring. Mutations that occur in

More information

CCR Biology - Chapter 9 Practice Test - Summer 2012

CCR Biology - Chapter 9 Practice Test - Summer 2012 Name: Class: Date: CCR Biology - Chapter 9 Practice Test - Summer 2012 Multiple Choice Identify the choice that best completes the statement or answers the question. 1. Genetic engineering is possible

More information

MS. Structure, Function, and Information Processing

MS. Structure, Function, and Information Processing MIDDLE SCHOOL LIFE SCIENCE Alignment with National Science Standards Use the chart below to find Science A-Z units that best support the Next Generation Science Standards* for Middle School Life Science,

More information

11.1. A population shares a common gene pool. The Evolution of Populations CHAPTER 11. Fill in the concept map below.

11.1. A population shares a common gene pool. The Evolution of Populations CHAPTER 11. Fill in the concept map below. 11.1 GENETIC VARIATION WITHIN POPULATIONS Study Guide KEY CONCEPT A population shares a common gene pool. VOCABULARY gene pool allele frequency MAIN IDEA: Genetic variation in a population increases the

More information

Algorithms in Computational Biology (236522) spring 2007 Lecture #1

Algorithms in Computational Biology (236522) spring 2007 Lecture #1 Algorithms in Computational Biology (236522) spring 2007 Lecture #1 Lecturer: Shlomo Moran, Taub 639, tel 4363 Office hours: Tuesday 11:00-12:00/by appointment TA: Ilan Gronau, Taub 700, tel 4894 Office

More information

Smart Science Lessons and Middle School Next Generation Science Standards

Smart Science Lessons and Middle School Next Generation Science Standards Smart Science Lessons and Middle School Next Generation Science Standards You have chosen the right place to find great science learning and, beyond learning, how to think. The NGSS emphasize thinking

More information

1. When new cells are formed through the process of mitosis, the number of chromosomes in the new cells

1. When new cells are formed through the process of mitosis, the number of chromosomes in the new cells Cell Growth and Reproduction 1. When new cells are formed through the process of mitosis, the number of chromosomes in the new cells A. is half of that of the parent cell. B. remains the same as in the

More information

Grade Stand Sub-Strand Standard Benchmark GRADE 6

Grade Stand Sub-Strand Standard Benchmark GRADE 6 Grade Stand Sub-Strand Standard Benchmark OF OF OF A. Scientific World View B. Scientific Inquiry C. Scientific Enterprise understand that science is a way of knowing about the world that is characterized

More information

High School Science Course Correlations between Ohio s 2010 Course Syllabi and the First Draft of the High School NGSS

High School Science Course Correlations between Ohio s 2010 Course Syllabi and the First Draft of the High School NGSS High School Science Course Correlations between Ohio s 2010 Course Syllabi and the First Draft of the High School NGSS This document correlates the content in Ohio s course syllabi with the performance

More information

Unit 2 Metabolism and Survival Summary

Unit 2 Metabolism and Survival Summary Unit 2 Metabolism and Survival Summary 1 Metabolism pathways and their control (a) Introduction to metabolic pathways This involves integrated and controlled pathways of enzymecatalysed reactions within

More information

Section Review 15-1 1.

Section Review 15-1 1. Section Review 15-1 1. Beagle 2. theory of evolution 3. varied 4. Darwin s curiosity might have led him to make many observations and ask questions about the natural world. His analytical nature may have

More information

THE INTERESTING HISTORY OF CELLS STUDENT HANDOUT. There are two basic types of cells, prokaryotic cells and eukaryotic cells.

THE INTERESTING HISTORY OF CELLS STUDENT HANDOUT. There are two basic types of cells, prokaryotic cells and eukaryotic cells. THE INTERESTING HISTORY OF CELLS STUDENT HANDOUT There are two basic types of cells, prokaryotic cells and eukaryotic cells. Figure 1. A PROKARYOTIC CELL The prokaryotes are very small singlecelled organisms

More information

The Origin of Life. The Origin of Life. Reconstructing the history of life: What features define living systems?

The Origin of Life. The Origin of Life. Reconstructing the history of life: What features define living systems? The Origin of Life I. Introduction: What is life? II. The Primitive Earth III. Evidence of Life s Beginning on Earth A. Fossil Record: a point in time B. Requirements for Chemical and Cellular Evolution:

More information

FAQs: Gene drives - - What is a gene drive?

FAQs: Gene drives - - What is a gene drive? FAQs: Gene drives - - What is a gene drive? During normal sexual reproduction, each of the two versions of a given gene has a 50 percent chance of being inherited by a particular offspring (Fig 1A). Gene

More information

Campbell Biology in Focus Correlation for AP Biology Curriculum Framework

Campbell Biology in Focus Correlation for AP Biology Curriculum Framework Campbell Biology in Focus Correlation for AP Biology Curriculum Framework Chapters/ Graphical analysis of allele frequencies in a population 5 Application of the Hardy-Weinberg equilibrium equation 1,

More information

Chapter 13: Meiosis and Sexual Life Cycles

Chapter 13: Meiosis and Sexual Life Cycles Name Period Chapter 13: Meiosis and Sexual Life Cycles Concept 13.1 Offspring acquire genes from parents by inheriting chromosomes 1. Let s begin with a review of several terms that you may already know.

More information

Given these characteristics of life, which of the following objects is considered a living organism? W. X. Y. Z.

Given these characteristics of life, which of the following objects is considered a living organism? W. X. Y. Z. Cell Structure and Organization 1. All living things must possess certain characteristics. They are all composed of one or more cells. They can grow, reproduce, and pass their genes on to their offspring.

More information

Macroevolution: Change above the species level NABT 2006 Evolution Symposium

Macroevolution: Change above the species level NABT 2006 Evolution Symposium Introduction Macroevolution: Change above the species level NABT 2006 Evolution Symposium The basic concept of evolution change over time can be examined in two different time frames. The first, which

More information

Scottish Qualifications Authority

Scottish Qualifications Authority National Unit specification: general information Unit code: FH2G 12 Superclass: RH Publication date: March 2011 Source: Scottish Qualifications Authority Version: 01 Summary This Unit is a mandatory Unit

More information

Lecture 3: Mutations

Lecture 3: Mutations Lecture 3: Mutations Recall that the flow of information within a cell involves the transcription of DNA to mrna and the translation of mrna to protein. Recall also, that the flow of information between

More information

Cell Division. Use Target Reading Skills. This section explains how cells grow and divide.

Cell Division. Use Target Reading Skills. This section explains how cells grow and divide. Cell Processes and Energy Name Date Class Cell Processes and Energy Guided Reading and Study Cell Division This section explains how cells grow and divide. Use Target Reading Skills As you read, make a

More information

Key Questions. How is evolution defined in genetic terms?

Key Questions. How is evolution defined in genetic terms? Getting Started Objectives 17.1.1 Define evolution in genetic terms. 17.1.2 Identify the main sources of genetic variation in a population. 17.1.3 State what determines the number of phenotypes for a trait.

More information

Basic Biological Principles Module A Anchor 1

Basic Biological Principles Module A Anchor 1 Basic Biological Principles Module A Anchor 1 Key Concepts: - Living things are made of units called cells, are based on a universal genetic code, obtain and use materials and energy, grow and develop,

More information