Table of Contents Overview and Introduction: ................................................................... 8 1: Overview of this course, and the instructional sequence presented. ......................................... 8 2: The AP exam. .................................................................................................................................... 10 A Tour of the Science Practices ....................................................... 15 SP 1: Use Representations and Models to Communicate Scientific Phenomena and Solve Scientific Problems................................................................................................................................. 15 1. SP 1 discussion ............................................................................................................................ 15 SP 2. Use Mathematics Appropriately. ............................................................................................. 16 1. SP 2 Discussion ............................................................................................................................. 16 2. MATH Skills: Metric system......................................................................................................... 17 3. MATH Skills: Dilutions .................................................................................................................. 19 SP 3. Engage in scientific questioning to extend thinking or to guide investigations within the course. .............................................................................................................................................. 20 1. SP 3 Discussion ............................................................................................................................. 20 SP 4. Plan and implement data collection strategies appropriate to a particular scientific question. .................................................................................................................................................. 21 1. SP 4 Discussion ........................................................................................................................... 21 SP 5. Perform Data Analysis and Evaluation of Evidence. .......................................................... 22 1. SP 5 Discussion ........................................................................................................................... 22 2. MATH Skills: Descriptive statistics .............................................................................................. 23 3. MATH Skills: Standard Deviation ................................................................................................ 25 4. MATH Skills: Standard Error ....................................................................................................... 27 SP 6. Work with scientific explanations and theories. ................................................................... 34 1. SP 6 Discussion ............................................................................................................................. 34 SP 7. Connect and relate knowledge across various scales, concepts, and domains. ........... 35 1. SP 7 Discussion ............................................................................................................................. 35 Domain 1: Evolution ........................................................................... 37 1.1: Natural selection is a major mechanism of evolution. (EK1.A.1) ........................................... 37 1. Natural Selection ........................................................................................................................... 37 2. The Modern Synthesis ................................................................................................................. 38 1.2: Natural selection acts on phenotypic variations in populations. (EK1.A.2) ......................... 40 1. How Natural Selection Works...................................................................................................... 40 1 1.3: Evolutionary change is also driven by random processes. (EK1.A.3) ................................. 42 1. Other Evolutionary Forces ........................................................................................................... 42 1.4: Biological evolution is supported by scientific evidence from many disciplines, including mathematics. (EK1.A.4) ........................................................................................................................ 45 1. Evidence of Evolution .................................................................................................................... 45 2. MATH Skills: HW Theory ............................................................................................................. 47 1.5: Organisms share many conserved core processes and features that evolved and are widely distributed among organisms today. (EK1.B.1) ................................................................... 50 1. Evidence of Common Ancestry ................................................................................................... 50 1.6: Phylogenetic trees and cladograms are graphical representations (models) of evolutionary history that can be tested. (EK1.B.2) ................................................................................................ 51 1. Phylogeny ...................................................................................................................................... 51 1.7: Speciation and extinction have occurred throughout the Earth’s history. (EK1.C.1) ...... 54 1. Speciation Concepts ....................................................................................................................... 54 1.8: Speciation may occur when two populations become reproductively isolated from each other. (EK1.C.2) .................................................................................................................................... 55 1. Speciation process.......................................................................................................................... 56 1.9: Populations of organisms continue to evolve. (EK1.C.3) ....................................................... 57 1. Ongoing evolution of organisms ................................................................................................... 57 1.10: There are several hypotheses about the natural origin of life on Earth, each with supporting scientific evidence. (EK1.D.1).......................................................................................... 58 1. Origin of life .................................................................................................................................... 58 Domain 2: Matter ................................................................................ 63 2.1: Organisms must exchange matter with the environment to grow, reproduce and maintain organization. (EK2.A.3)....................................................................................................................... 63 1. Matter exchange ............................................................................................................................. 63 2. Properties of Water ........................................................................................................................ 65 3. MATH Skills: pH ........................................................................................................................... 66 4. Constraints on Cell Size ................................................................................................................ 67 5. MATH Skills: Surface Area:Volume Ratio .................................................................................. 69 2.2: The subcomponents of biological molecules and their sequence determine the properties of that molecule. (EK4.A.1) ................................................................................................................. 70 1. Biological Molecules ..................................................................................................................... 70 2.3: Variation in molecular units provides cells with a wider range of functions. (EK4.C.1) .. 71 1. Variation in Biological Molecules- Carbohydrates and Lipids .................................................. 72 2. Variation in Biological Molecules- Proteins .................................................................................. 72 2 3. Variation in Biological Molecules- Nucleic Acids......................................................................... 74 2.4: Cell membranes are selectively permeable due to their structure. (EK2.B.1) .................. 75 1. Cell Membrane Structure ............................................................................................................... 75 2.5: Growth and dynamic homeostasis are maintained by the constant movement of molecules across membranes. (EK2.B.2)........................................................................................ 77 1. Mechanisms of Cellular Transport .............................................................................................. 77 2. Analyzing Transport ........................................................................................................................ 79 3. MATH Skills: Water Potential and Solute Potential .................................................................. 81 2.6: Eukaryotic cells maintain internal membranes that partition the cell into specialized regions. (EK2.B.3) ............................................................................................................................... 83 1. Cellular Compartmentalization ....................................................................................................... 84 2. Major Eukaryotic Organelles .......................................................................................................... 85 2.7: The structure and function of subcellular components, and their interactions, provide essential cellular processes. (EK4.A.2) ............................................................................................. 86 1. Organelle structure and function- information processing .......................................................... 87 2. Organelle structure and function- matter and energy processing............................................ 87 Domain 3: Energy ............................................................................... 90 3.1: All living systems require constant input of free energy. (EK2.A.1) .................................... 90 1. Bioenergetic Theory ...................................................................................................................... 90 2. MATH Skills: Gibbs Free Energy ................................................................................................ 91 3. Metabolic Strategies ....................................................................................................................... 94 4. MATH Skills: Coefficient Q10 ...................................................................................................... 96 3.2: Interactions between molecules affect their structure and function. (EK4.B.1) .................. 98 1. Enzyme structure and function .................................................................................................... 98 2. Regulation of Enzyme Activity..................................................................................................... 100 3.3: Organisms capture and store free energy for use in biological processes. (EK2.A.2) . 102 1. Energy Processing ...................................................................................................................... 102 2. Photoautotrophic nutrition- light reactions ................................................................................. 104 3. Photoautotrophic nutrition 2- Carbon fixation ........................................................................... 106 4. Chemoheterotrophic nutrition- anaerobic cellular respiration .................................................. 107 5. Chemoheterotrophic nutrition: Aerobic cellular respiration .................................................... 109 3.4: Cooperative interactions within organisms promote efficiency in the use of energy and matter. (EK4.B.2) ................................................................................................................................ 111 1. Organizational efficiency in energy processing. ....................................................................... 111 Domain 4: Information ...................................................................... 114 3 4.1: DNA, and in some cases RNA, is the primary source of heritable information. (EK3.A.1) ................................................................................................................................................................ 114 1. DNA Structure ............................................................................................................................. 114 2. DNA Replication............................................................................................................................ 117 3. Protein Synthesis ........................................................................................................................ 119 4. Genetic Information Processing & Expression ......................................................................... 121 5. Genetic Engineering- Techniques ............................................................................................... 123 6. Genetic Engineering- Applications............................................................................................... 125 4.2: In eukaryotes, heritable information is passed to the next generation via processes that include the cell cycle and mitosis or meiosis plus fertilization. (EK3.A.2) ............................... 128 1. Mitosis .......................................................................................................................................... 132 2. Cell Cycle Control....................................................................................................................... 133 3. Meiosis ........................................................................................................................................... 135 4.3: The chromosomal basis of inheritance provides an understanding of the pattern of passage (transmission) of genes from parent to offspring. (EK3.A.3) ....................................... 138 1. Mendelian Genetics .................................................................................................................... 138 2. MATH SKILLS: Genetics Probabilities ...................................................................................... 141 3. Chromosomal Disorders ............................................................................................................... 137 4.4: The inheritance pattern of many traits cannot be explained by simple Mendelian genetics. (EK3.A.4) ................................................................................................................................................ 143 1. Mendelian Extensions ................................................................................................................. 143 2. Non-Mendelian Traits .................................................................................................................. 143 4.5: Gene regulation results in differential gene expression, leading to cell specialization. (EK3.B.1) ................................................................................................................................................ 146 1. Gene Regulation ......................................................................................................................... 146 2. Prokaryotic Gene Regulation ..................................................................................................... 148 3. Eukaryotic Gene Regulation ...................................................................................................... 150 4. Gene Regulation Phenotype................................................................................................... 152 4.6: A variety of intercellular and intracellular signal transmissions mediate gene expression. (EK3.B.2) ................................................................................................................................................ 153 1. Signal Control of Gene Expression .......................................................................................... 153 4.7: Changes in genotype can result in changes in phenotype. (EK3.C.1).............................. 126 1. Mutations...................................................................................................................................... 126 4.8: Biological systems have multiple processes that increase genetic variation. (EK3.C.2) 145 1. Generation of Variation .............................................................................................................. 145 4 4.9: Viral replication results in genetic variation, and viral infection can introduce genetic variation into the hosts. (EK3.C.3)................................................................................................... 128 1. Viral Genetics................................................................................................................................ 128 4.10: Environmental factors influence the expression of the genotype in an organism. (EK4.C.2)................................................................................................................................................ 154 1. Environmental Effects on Phenotype. ....................................................................................... 154 Domain 5: Regulation....................................................................... 155 5.1: Timing and coordination of specific events are necessary for the normal development of an organism, and these events are regulated by a variety of mechanisms. (EK2.E.1) .......... 155 1. Development ................................................................................................................................ 155 5.2: Interactions between external stimuli and regulated gene expression result in specialization of cells, tissues and organs. (EK4.A.3) ................................................................. 157 1. Differentiation ............................................................................................................................... 157 5.3: Organisms use feedback mechanisms to maintain their internal environments and respond to external environmental changes. (EK2.C.1) ............................................................. 158 1. Feedback Loops.......................................................................................................................... 158 5.4: Homeostatic mechanisms reflect both common ancestry and divergence due to adaptation in different environments. (EK2.D.2) ............................................................................ 160 Discussion 1: ..................................................................................................................................... 160 Discussion 2: ..................................................................................................................................... 162 5.5: Biological systems are affected by disruptions to their dynamic homeostasis. (EK2.D.3) ................................................................................................................................................................ 162 1. Effects of Disruptions ................................................................................................................. 162 5.6: Plants and animals have a variety of chemical defenses against infections that affect dynamic homeostasis. (EK2.D.4) ...................................................................................................... 164 1. Immune Systems. ....................................................................................................................... 165 5.7: Timing and coordination of behavior are regulated by various mechanisms and are important in natural selection. (EK2.E.2) ........................................................................................ 167 Discussion 1: ..................................................................................................................................... 167 Domain 6: Communication .............................................................. 170 6.1: Cell communication processes share common features that reflect a shared evolutionary history. (EK3.D.1) ................................................................................................................................ 170 1. Introduction to Communication .................................................................................................. 170 6.2: Cells communicate with each other through direct contact with other cells or from a distance via chemical signaling. (EK3.D.2) .................................................................................... 171 5 1. Types of Cellular Signals ............................................................................................................ 171 6.3: Signal transduction pathways link signal reception with cellular response. (EK3.D.3) ... 173 1. Signal Transduction Pathways .................................................................................................... 173 6.4: Changes in signal transduction pathways can alter cellular response. (EK3.D.4) .......... 176 1. Alterations to Signaling Pathways............................................................................................... 176 6.5: Individuals can act on information and communicate it to others. (EK3.E.1) .................. 177 1. Communication Between Organisms......................................................................................... 178 6.6: Animals have nervous systems that detect external and internal signals, transmit and integrate information, and produce responses. (EK3.E.2) ............................................................ 180 1. Neurons ......................................................................................................................................... 180 2. Nervous Systems .......................................................................................................................... 184 Domain 7: Interactions ..................................................................... 186 7.1: Organisms exhibit complex properties due to interactions between their constituent parts. (EK4.A.4) ................................................................................................................................................ 186 1. Physiological Interactions ............................................................................................................. 186 7.2: Organisms respond to changes in their external environments. (EK2.C.2) ...................... 188 1. Responses to the Environment ................................................................................................. 188 7.3: Interactions among living systems and with their environment result in the movement of matter and energy. (EK4.A.6) ........................................................................................................... 189 1. Energy and Matter Acquisition .................................................................................................. 190 2: MATH SKILLS: Productivity Calculations ................................................................................ 193 7.4: All biological systems from cells and organisms to populations, communities and ecosystems are affected by complex biotic and abiotic interactions involving exchange of matter and free energy. (EK2.D.1) .................................................................................................. 194 1. Limiting Factors............................................................................................................................. 194 7.5: The level of variation in a population affects population dynamics. (EK4.C.3) ............... 196 1. Population Diversity ...................................................................................................................... 196 7.6: Interactions between and within populations influence patterns of species distribution and abundance. (EK4.B.3) ........................................................................................................................ 197 1. Community Interactions ................................................................................................................ 198 7.7: Communities are composed of populations of organisms that interact in complex ways. (EK4.A.5) ................................................................................................................................................ 199 1. Measuring Communities ............................................................................................................... 199 2. MATH SKILLS: Population Growth Equations ......................................................................... 200 7.8: The diversity of species within an ecosystem may influence the stability of the ecosystem. (EK4.C.4) ......................................................................................................................... 202 6 1. Ecosystem Stability ....................................................................................................................... 202 7.9: Distribution of local and global ecosystems changes over time. (EK4.B.4) ..................... 204 1. Ecosystem Changes ..................................................................................................................... 204 7 Overview and Introduction: 1: Overview of this course, and the instructional sequence presented. AP Biology is a survey of introductory college-level biology. Two major parts: The practice of being a biologist (the Science Practices), and the knowledge that biological investigations have generated about the functioning of living systems. We can’t spend a lot of time on the first part in video lessons. But the skills that you use can and should be applied to the second part as much as possible. Given the format, a lot of this application is going to have to happen by yourself. in the format. I’m here to help, but I can only do so much For you to get the most out of this process, you will need to take a very active role in your learning. You can sit and passively watch the videos and hope the content sticks in your brain, or you can actively engage with the material that we will be discussing, writing notes and jotting down the questions 8 that you have as you go through the videos. Then you can work to get the answers to those questions in a lot of different places (your teacher, your textbook, and the internet are three obvious sources). We will take a brief tour of the seven science practices first to make sure we are all speaking the same language, and then we will move in to the content of the course. I have organized the course content in to seven “domains”, each of which deal with one major aspect of living systems. I have a preferred order to move through them, but they should be able to be viewed in whatever order works best for you (though within a domain, it’s best to start at the beginning and move your way through sequentially). This organization may be somewhat different from the order of your class. That’s okay (it’s actually a good thing, because you’ll get a different way to look at the content of this course). The material that we will discuss is aligned to the AP Biology Curriculum Framework, which is published by the College Board, and freely available to anyone with an internet connection. You 9 can get a copy if you really want, but it’s written for teachers, and may not be so easy for students to use. Important note: The examples that I use are not the only examples that can be used to address the concepts of this course. I’ve marked the examples with the notation “Ex.”, so that you can easily identify them, and try to supply alternative examples if you can (that will be really helpful for you). 2: The AP exam. The AP Biology exam is a 3-hour exam, administered in mid-May. Depending on your school’s policies you either have to take it, or will be encouraged to take it (and you should take it). You are allowed to bring pencils, pens, and a four-function scientific calculator (a square root function is also allowed). A copy of the formula sheet will be provided for you (you can not use your own). The Curriculum Framework contains a list of “learner objectives”. These are the things that you will be 10 expected to do on the exam. If you consult the learner objectives, you will notice that they are written in such a way as to allow for a wide variety of questions. Demo Formula Sheet Sample Learner Objective: LO 1.1 The student is able to convert a data set from a table of numbers that reflect a change in the genetic makeup of a population over time and to apply mathematical methods and conceptual understandings to investigate the cause(s) and effect(s) of this change. [See SP 1.5, 2.2] The exam is broken into two sections: Section 1: Multiple Choice and Grid-In Questions: There are 63 multiple-choice questions and 6 mathematical grid-in questions. Each multiple choice question has four answer choices. Many of these questions will be on the content of the course directly, and many of the questions will analyze your ability to read and consider scientific information and apply that information to the 11 understandings of the course. The grid-in questions will require you to perform a particular mathematical analysis (including BUT NOT LIMITED to the equations on the formula sheet). will be administered over a period of 80 minutes. Section 1 Following section 1, you will have a 10-minute break period before moving on to section 2. Section 2: Constructed Response Questions: constructed response questions. There are 8 2 of these questions will be multi-part essays, and 6 of them will be smaller, single-part written responses. The constructed response questions will also assess your understanding of the course content and your ability to consider and apply scientific information to the study of Biology. At least one question will require you to graphically represent information, and your ability to design and analyze experiments will also be a fundamental feature of this section. To begin section 2, You will have a 10-minute reading period, followed by an 80 minute period in which to write your answers. Part 1 of the exam is graded by machine. Part 2 of the exam is graded at a “reading” which takes place in 12 mid-June, during which hundreds of AP Biology teachers and College Professors meet to read and score the questions. Exam grades are on a 1-5 scale. If you are planning on attending college, high scores (4 or 5) will often allow you to receive credit for introductory-level Biology courses, however the specific policies on college credit vary from institution to institution. Exam strategies: Answer every question: exam. There is no penalty for guessing on the You cannot get full credit in question two if you do not answer all parts of every question. Time yourself: For section one, a good rule of thumb is to spend no more than 1 minute per question. This will give you 12 minutes at the end to go back and work on any very difficult questions. In part 2, you should plan to spend about half of your time answering the first two, long essays, and then spend the remaining time addressing the final six. If you have spent 20 minutes a piece on the first two questions (40 minutes in total), you have ~6 minutes for each of the shorter questions. Write succinctly: make sure to answer the question you are being asked completely, but only write words that answer the question. 13 There are no points for literary elements in your writing. Opening and closing paragraphs are useless and waste time. Answer the question you are being asked: question. Do not stray from the Make sure to address the question and then move on. Get a good night’s sleep before the exam, and have a positive attitude: Both of these have been shown to have a significant effect on exam performance. 14 A Tour of the Science Practices SP 1: Use Representations and Models to Communicate Scientific Phenomena and Solve Scientific Problems. 1. SP 1 discussion Models are representations of thoughts. Models can be physical, computational, mathematical, or literally any other way that a thought can be represented (though in science, they tend to be one of the first three, or maybe verbal). You are expected to be able to create, describe, refine, use, and re-express representations and models in the work that you do in this course. Be careful: Do not make the mistake of confusing a model for the reality that it represents. Models are always simplifications of reality. 3 ways to work on developing your modeling/representation skills: When you are shown models or representations, take a moment and acknowledge that is what is happening. 15 Create alternate models and representations of the concepts that you are investigating from the ones that you are being shown by your teacher. Develop your own examples of phenomena under consideration. Take models and representations that you are using in class, and identify the simplifications in them. Refine the model to show this additional complexity (if possible—things can get very complicated very quickly). SP 2. Use Mathematics Appropriately. 1. SP 2 Discussion Math is the most objective way we have to describe and model scientific phenomena. Math is as important to biology as any other tool. You are expected to be able to justify the selection of a mathematical routine, apply a mathematical routine, and estimate numerical quantities. Biologists have an unfair reputation for not liking math. acceptable. This is not Math must be embraced, and loved. 16 3 ways to work on developing your math skills: Practice, practice, practice. Keep trying to use the math of this course wherever possible. Try to come up with estimates of ridiculously big numbers. How many hairs are on your head? body? How many cells are in your The point is not to get the right answer, but to work through the process of estimating such things. When you see math being used in class, try to justify why the math is appropriate for the task it is being used for. 2. MATH Skills: Metric system The metric system is the measurement system of science. It is based on powers of 10, which modify a base quantity (assigned as 100, or as we all know it, 1) of the major dimensions of matter, energy, and time in the universe. Prefixes are used to denote every power of 10 from 10-3 through 103, and then every three powers of ten larger and smaller. Biology ranges all over the metric system. 17 Selected prefixes and their quantities are provided on the formula sheet that you will have during the AP exam. Factor Prefix Symbol 109 giga G 106 mega M 103 kilo k 10-2 centi c 10-3 milli m 10-6 micro μ 10-9 nano n 10-12 pico p There are a lot of benefits to using the metric system. The common powers of 10 structure makes arithmetic with widely different numbers very easy. It is universal among all scientists on the planet. A kilogram always signifies the same mass. 18 You need to use the metric system in all data recording and data calculations that you engage in during this course. 3. MATH Skills: Dilutions The dilution equation is provided on the formula sheet that you will have during the AP exam. It is useful in lab. You may use it or not, but you need to be able to use it (for the exam, and in your life as a competent laboratory scientist: C = concentration V = Volume i = initial f = final 19 You will know (or be able to figure out) three of these variables. You will use the equation to figure out the fourth. Sample Problem: How much of a solution of 2M sucrose must be diluted to make 3L of a 1M sucrose solution? ANSWER: 1.5 L SP 3. Engage in scientific questioning to extend thinking or to guide investigations within the course. 1. SP 3 Discussion Not all questions can be addressed scientifically. There are inherent limitations on the kinds of questions that science can answer. Example: What happens to us when we die? Inherently limited in what science can tell us Scientific answer: our bodies are decomposed back in to simpler compounds, which are reincorporated by the ecosystems of the planet. 20 You need to be able to pose, refine, and evaluate scientific questions. 3 ways to work on developing your scientific questioning skills: Question how the knowledge that you are learning in this course was determined. Work on asking scientifically useful/interesting questions…and then refining them in to even better questions. Analyze when a question is not going to be appropriate for scientific investigation. SP 4. Plan and implement data collection strategies appropriate to a particular scientific question. 1. SP 4 Discussion Science relies on data to help answer questions. Data is generated in many different circumstances, including field investigations and experimentation. The circumstances in which data is collected will influence how useful that data is for answering questions. 21 You need to be able to justify the selection of data, design a plan to collect data, collect data, and evaluate sources of data. 3 ways to work on developing your data collection skills: Identify which data sources will be most useful for answering particular scientific questions. Determine what sorts of equipment and procedures are used to collect specific types of data. Use the data sets generated by other scientists to develop your answers to scientific questions. SP 5. Perform Data Analysis and Evaluation of Evidence. 1. SP 5 Discussion The analysis of data is how science develops explanations. Analysis and evaluation of data include statistical, graphical, and computational manipulation of a data source. 22 You need to be able to analyze data, refine observations and measurements based on the analysis of data, and evaluate the evidence provided by a data set. 3 ways to work on developing your data analysis skills: When investigating a particular scientific explanation, evaluate the data that supports (and refutes) the explanation. In experimental settings, use the data that is being generated to determine if the data is appropriate for the experiment. Based on this analysis, refine your experimental measurements if necessary. Determine the statistical aspects of a particular data set, and consider how they support or refute a particular explanation for the data set. 2. MATH Skills: Descriptive statistics Descriptive statistics are used to tell us about the characteristics of a particular data set. 23 Each descriptive statistic tells us something different, and may be more or less useful depending on the data set, and the concepts it is supporting or refuting. Mean, median, mode, range Sample data set A: Mean: 5cm, 5 cm, 10 cm, 20 cm The sum of all of the data points in a data set, divided by the number of items in the data set. Explained on the Formula Sheet verbally and also presented in sigma (sum) notation: _ x = mean n = size of the sample (number of items) Mean of sample data set A: 10cm 24 Mode: The most common value among the items in the data set. Mode of sample data set A: 5 cm Median: The middle value of the data set, or the mean of the middle two values in the data set. Median of sample data set A: Range: 7.5 cm The difference between the highest value and lowest value in the data set. Range of sample data set: 3. MATH Skills: Standard Deviation: the data set. 15 cm Standard Deviation A measurement of the variance of the items in “How far away from the mean are the items in the data set?” 25 The higher the standard deviation, the further away most data points are from the mean of the data set. The equation is on the formula sheet in sigma notation. Almost all data in a data set will lie within +/- 3 standard deviations of the mean if the data has a normal distribution (“bell curve”). 68% of the data will lie within 1 standard deviation, 95 % will be within two standard deviations, and 99.7% will be within three standard deviations (if the data has a normal distribution). s = standard deviation Sample data set A: std. dev: Sample data set B: std. dev: 5 cm, 5 cm, 10 cm, 20 cm Mean: 10 cm Mean: 10 cm ~7 8 cm, 9 cm, 11 cm, 12 cm ~2 26 These two data sets have the same mean, but their variance is different. The meaning of this depends on what the data refers to. 4. MATH Skills: Standard Error: Standard Error A measurement of the variation in the means of data sets taken from the same population. usually have different means. those means will be. Different samples will Standard error tells us how varied It is the standard deviation of the means in a sample set. Useful because if two data sets have means that are more than 2 standard errors away from each other (actually 1.96) have a 95% confidence that they are statistically significant. This is typically the cut-off for being able to reject a null hypothesis (that the variance between our observation and our expectation is due to chance) 27 SE = standard error Standard error of sample data set A: ~3.5 Standard error of sample data set B: ~1 Not a particularly useful statistic for this data set, since the means of both samples are the same to begin with. Two different data sets: Sample set 1: Mean: ~6.6g 6.2 g, 6.4 g, 6.6 g, 7.0 g stdev= +/- ~0.34g st. error: +/- ~.17 Sample set 2: Mean = ~12.5g 12.2 g, 12.3 g, 12.6 g, 13.0 g stdev= +/- ~0.359g st. error: +/- ~.18 Here, there is a difference in the means, the standard deviations, and the standard error of the two data sets. 28 The non-overlap indicates that there is at least a 95% confidence that the differences between these two means is not due to chance fluctuations in our data set 5. MATH Skills: Hypothesis Testing Our statistical expectation is NOT the same thing as our experimental expectation. The hypothesis of the experiment governs how we frame our statistical expectation, but our statistical hypothesis is ALWAYS the null hypothesis (that the variation between expectation and observation is due to chance). Chi-square: A way to determine if the variance between what we observe and what we expect in a set of categorical data is statistically significant or not. X2= chi-square value o = observed values e = expected values 29 It works best to calculate chi-square data methodically, using a table approach. The major areas where categorical data will be encountered in this course will be in ecology (ex. distribution of organisms in an environment) and mendelian genetics (ex. Number of progeny that have certain inherited characteristics). Sample problem: An ecologist is habitat preferences of periwinkles on the rocky coast line of the New England coast. She hypothesizes that more periwinkles will be found closer to the tide line. To test her hypothesis, she collects data by counting the number of periwinkles within a .5 m2 quadrat sample that she observes on a rocky coast line location at low tide: Distance from low tide: Number of periwinkles observed: At low tide line: 1 meter above low tide: 36 24 30 2 meters above low tide: 10 3 meters above low tide: 3 4 meters above low tide: 2 Total: 75 Determine if the difference in the number of periwinkles observed in each location is statistically significant: Step 1: Frame the null hypothesis- “The difference between the number of periwinkles that we observe at each location and the number that we expect is due to chance.” Fine, but what do we expect? This step is a bit tricky. Given the experimental hypothesis, you might want to state that we expect more periwinkles closer to the tide line. But that would make it impossible to do a chi-square analysis on this data (how many of the periwinkles should we include for “more”?). So, our statistical expectation is going to be different from our experimental expectation. For this analysis, the statistical expectation is that periwinkles have no location preference. This way, if the null hypothesis is rejected, then we are saying that there is a 31 statistically significant difference in the distribution of periwinkles on the shoreline, which supports our experimental hypothesis. Also, by setting up our statistical test like this, we can easily determine the “expected” values for each of our categories (equal numbers of the total periwinkles that we have observed). Chi Square Determination: Category: o(observed) e(expected) o-e o-e2 (o-e2)/e At low tide 36 15 21 441 29.4 1 meter above 24 15 9 81 5.4 2 meters above 10 15 -5 25 1.7 3 meters above 3 15 -12 144 4 meters above 2 15 -13 169 X2 = 9.6 11.2 57.3 32 How to use the X2 value: We compare the chi-square value to a table of values, according to the number of degrees of freedom in the data set (the number of categories – 1): A p value less than or equal to .05 is the typical “cut-off” for rejecting the null hypothesis (ie. There must be at least a 95% probability that the variance we observe from what we expect is NOT due to chance in order for us to reject the null hypothesis). Our answer from above has 5 degrees of freedom (5 categories of data). Comparing our value of 57.3 to the chi-square table, we can see that our value is above the .05 cutoff of 9.49 (it’s actually well above the .01 cutoff). This means that our null hypothesis is rejected, and an alternative hypothesis (that the difference between the observed and expected value is not due to chance) is not rejected. The results of this statistical hypothesis test along with the observed data support our experimental hypothesis. 33 If we don’t reject the null, it DOES NOT mean that the null is accepted. It just means that it has failed to be rejected. SP 6. Work with scientific explanations and theories. 1. SP 6 Discussion Scientific explanations are always tentative, and supported by evidence. The nature of science is not about “proving” anything. Science works by falsifying possible explanations (“hypotheses”). A scientific explanation must be testable. If an explanation cannot be tested, science can not support or refute it. Scientific explanations that support a broad range of observable phenomena are called “theories”. Theories can be refined over time to account for additional observations, or they can be refuted, but they can never be “proven”. You need to be able to justify claims with evidence, construct explanations based on evidence, articulate the reasons an 34 explanation or theory is refined or replaced, and evaluate alternative scientific explanations. 3 ways to develop your scientific explanation skills: Never make a claim without evidence to support it (remember that not all evidence is created equal). Consider the evidence that supports the explanations and theories that you will be exposed to in this course. Explain how and why they have had to be revised over time. Consider alternative explanations for the phenomena that you investigate in this course. Analyze how the available evidence supports or refutes these alternatives and determine possible evidence that would support or refute them SP 7. Connect and relate knowledge across various scales, concepts, and domains. 1. SP 7 Discussion Science connects concepts that span several spatial and temporal scales (in biology, phenomena move from the atomic level to the entire biosphere, and from fractions of a second to billions of years). 35 Science also depends on many different kinds of information from a variety of domains to make explanations. Biological explanations use physical, chemical, and mathematical information. You need to be able to connect phenomena and models across spatial and temporal scales, and connect concepts across domains to generalize and extrapolate ideas. 3 ways to develop your “scale-domain” thinking: Actively consider how the concepts of this course rely on knowledge from a variety of domains. Spend time relating temporal scales and spatial scales to the ones that you are most familiar with. Consider how biological explanations agree or conflict with explanations from other sciences. 36 Domain 1: Evolution 1.1: Natural selection is a major mechanism of evolution. (EK1.A.1) 1. Natural Selection Darwin/Historical Evolutionary Development Historical development of evolutionary thought pre-Darwin Lyell- Geology, Uniformatarianism very old earth. Malthus- Exponential Population Growth LaMarck- Evolution. Inheritance of acquired characteristics (wrong, but still evolutionary) Darwin- Uninspired youth. Ship’s naturalist on board HMS Beagle. Circumnavigation of the globe. Return to Europe and delay publication of theory until pressed by Alfred Russell Wallace. Natural Selection: Inherent Variation in all organisms. Overproduction of Offspring (Malthus) 37 Differential success (“fitness”) of different variants in survival and reproduction (“adaptations”). Inheritance of adaptations leads to populations becoming better adapted for the environment over time. Repeat for millions of years. Fundamental conclusions: Ancient age of the Earth (billions of years, not thousands) Common Ancestry of all organisms on Earth (tree thinking). Unsettled by Darwin: Origin of Life Origin of species Nature of variation/inheritance 2. The Modern Synthesis Connection of evolution to change in allele frequencies in populations/ basic mathematical analysis. The “Modern Synthesis”- connecting Darwinian evolution to genetics and modern understanding of inheritance. 38 Traits are the result of genetic instructions in DNA (“genes”) Variations in traits are the result of changes to genes (“mutations”) “alleles”: different variants in traits Evolution: Example: Changes in allele frequencies over time. Galapagos Finches/Grant Studies Studies of finch populations on an isolated island in the Galapagos. Measurements of the beak dimensions of all birds on the island every year for decades. Able to connect changes in beak dimensions to fluctuations in the environment (precipitation, seed sizes) Relevant Misconceptions: Evolution is a “population level” phenomenon. The evolution of a population emerges from the individual fitness of members of that population. As they survive and reproduce, or not, the frequencies of alleles in the next generation will change accordingly. 39 1.2: Natural selection acts on phenotypic variations in populations. (EK1.A.2) 1. How Natural Selection Works. Connection of Phenotype to Genotype (Basic Central Dogma) Genotype: The alleles that an individual has for a particular trait. Homozygous: Heterozygous: Two copies of the same allele. Two copies of different alleles for each trait. Why do we have two alleles for each trait? Because of sexual reproduction. Phenotype: The trait that an individual shows. Genotype determines phenotype. Alleles control the production of proteins and the proteins determine traits Dominant vs. recessive alleles- some alleles (“dominant”) will control phenotype over other alleles (“recessive”) when both are present. Example: Eye color (simplified) 40 Two alleles: Brown (B) and blue (b). eyes and blue eyes. Two phenotypes: Brown Individuals who are homozygous will either have brown (BB) or blue (bb) eyes. Brown is dominant and blue is recessive, so heterozygotes will have brown eyes. Relevant Misconception: Dominant alleles are “better” than recessive alleles: Dominant and recessive have nothing to do with their effect on fitness. They only refer to how they contribute to phenotype expression. Relationship between phenotype and fitness: Different phenotypes will be more or less fit, depending upon the requirements of the environment. “Fitness”: Ability to contribute genes to the next generation (reproduction). The environment determines fitness, and fitness may change as the environment changes. Example: Pesticide Resistance 41 Individuals who are resistant to a pesticide will be more fit than individuals who are not when the pesticide is present in the environment. If there is no pesticide present, resistance will not contribute to fitness (may detract, depending on the nature of that resistance). Human impact on variation: Humans are able to impact variation in other organisms by controlling which individuals are able to reproduce. Artificial selection: When reproductive success is determined by human requirements Examples: Dogs, monocrop fields 1.3: Evolutionary change is also driven by random processes. (EK1.A.3) 1. Other Evolutionary Forces Genetic Drift: The random, non-selective, changes in allele frequency due to chance events. 42 Genetic drift has a larger effect on smaller populations, since each individual is a larger percentage of the total alleles in the population. The Founder Effect: When the descendants of a small, founding population have different percentages of alleles than the population that the founders came from. Example: The rate of polydactyly in the Amish population is higher than it is in the European population that the Amish came from. The Bottleneck Effect: Following a catastrophic decrease in a population, the survivors of the “bottleneck” event may have a different percentage of alleles than the pre-bottleneck population. Example: The Cheetah population underwent a major bottleneck during the last ice age (and subsequent hunting). Any two cheetahs are genetically equivalent to identical twins. 43 Gene Flow: The intermixing of alleles from two overlapping populations. This frequently leads to an “equalizing” of differences in allele frequencies between populations. Example: The effect of human migration on population characteristics since the industrial revolution. Sexual Selection: Selection for characteristics that aid an organism’s reproductive success. Often these characteristics may seem “maladaptive” for the individual. Examples: Peacocks (intersexual), dominance competitions (intrasexual) Relevant Misconceptions: Evolution is a “random” process: frequencies over time. Evolution is a change in allele There are random evolutionary processes (genetic drift) and non-random, selective processes (natural selection). 44 1.4: Biological evolution is supported by scientific evidence from many disciplines, including mathematics. (EK1.A.4) 1. Evidence of Evolution Geographical/Geological Evidence: Radiometric dating enables the age of fossils and rock layers to be determined and establishes the age of the Earth as ~4.5 billion years. The appearance of similar fossils on different continents correlates to the positions of those continents over geologic time, accounting for continental drift. The fossil record establishes a history of life on Earth. are no “anachronisms” in the fossil record. There Fish show up millions of years before mammals. Living organisms resemble fossilized, extinct forms. Fossilized organisms showing major evolutionary transitions between groups are found with regularity. Example: Tiktallik 45 Anatomical Evidence: Similarities and differences in the anatomy (morphology) of organisms provide evidence/clues to how they have evolved. Vestigial structures: Structures that have lost their primary adaptive purpose. Example: Whale hind limbs Homologous structures: Structures present in a common ancestor, which have diverged during evolution. Example: Vertebrate limbs. Analogous structures: Structures that have evolved multiple times in different lineages to fill similar adaptive needs. Example: Wings Chemical Evidence: Similarities and differences in DNA and protein sequences. As lineages diverge, they will accumulate mutations in DNA sequences, which will alter the sequences of proteins. 46 Mathematical Modeling: Computational analysis of evolution: The ability to analyze large amounts of chemical sequence data to establish evolutionary relationships among organisms. Hardy-Weinberg Theory: The ability to quantify the amount of evolutionary change from generation to generation to determine how evolution is affecting the population. 2. MATH Skills: HW Theory Hardy-Weinberg equations enable us to determine how much a population is evolving from generation to generation. “Hardy-Weinberg equilibrium”: population. Refers to an idealized, non-evolving Five characteristics: Large size (no genetic drift) Random mating (no sexual selection) Stable environment (no natural selection) No immigration/emigration (no gene flow) 47 No mutations. No real population is in HW equilibrium. Two equations (for a trait controlled by two alleles, where p is the dominant allele and q is the recessive allele): Gene pool equation: p + q = 1 p = frequency of dominant allele, q= frequency of recessive allele Organism equation: p2 + 2pq + q2 = 1 p2 = frequency of homozygous dominant individuals, 2pq = frequency of heterozygotes, and q2 = frequency of homozygous recessive individuals To solve HW equations, always determine the frequency of recessive individuals first, and use that to solve the rest of the equation. Sample problem: In pea plants, the allele for purple flowers is dominant to the allele for white 48 flowers. If 99% of the plants in the population have purple flowers, determine the percentage of heterozygotes in the population. Solution: Given: 99% (.99 frequency) have purple flowers. Can’t say how much are p2 and how much are 2pq. But we do know that p2 + 2pq + q2 = 1, and that in this case p2 + 2pq = .99. So we can solve for q2, which is .01 (1 percent). Now we solve for q, by taking the square root of .01 which is .1 (square root weirdness in decimal land!). percent). If q is .1, than p = 1 - .1, or .9 (90 Now that we know that, we can solve for the frequency of p2 which is .92 = .81, and the frequency of 2pq, which is 2*.9*.1 = .18. So 18% of our population are heterozygous for flower color. If the data in a HW problem does not add up to 1, then the population is NOT in HW equilibrium. Uses of HW equilibrium: To determine how a population is evolving from generation to generation. (Is the population out of HW equilibrium? Are the p and q frequencies changing over generations?) 49 To help to determine which evolutionary pressures are affecting a population more/less. 1.5: Organisms share many conserved core processes and features that evolved and are widely distributed among organisms today. (EK1.B.1) 1. Evidence of Common Ancestry Genetic Code: All cellular organisms use DNA to store genetic information, expressing that information through RNA in to protein structure. The “genetic code” that translates nucleic acid into protein structure is essentially universal among all lineages of life. Metabolic Pathways: All cellular organisms produce usable energy through similar metabolic pathways. Example: glycolysis is the first energy producing metabolic pathway in essentially every cell on the planet. Cellular Morphological similarities: 50 All cells on the planet are structured on one of two major organizational plans. Prokaryotes: Cells that lack any internal membrane-enclosed compartments (“organelles”) Eukaryotes: Cells that contain a variety of internal organelles. Endosymbiosis: The theory that major eukaryotic organelles (mitochondria and chloroplasts) evolved from free-living prokaryotic cells that were engulfed and internalized by eukaryotic ancestors. Evidence: DNA sequences, reproduction patterns, double membrane structure. 1.6: Phylogenetic trees and cladograms are graphical representations (models) of evolutionary history that can be tested. (EK1.B.2) 1. Phylogeny Phylogenetic tree/cladogram construction: A cladogram is a diagram that groups items according to the number of traits that they have in common: 51 The number of characteristics that are shared among the items are determined. The items are then arranged in a “tree diagram” to represent these similarities. A phylogenetic tree is a cladogram that groups organisms according to the number of evolutionary traits that they have in common. Characters/Sequence Data: To construct a phylogenetic tree, either “shared derived characters” are used (traits that are representative of the evolution of the organisms), or the amount of similarity in DNA/protein sequence information is looked at. Examples: vertebrate tree, whole life tree. Phylogenetic tree construction: Determine similarities among organisms (character table works well). Arrange organisms in a tree diagram showing simplest possible evolution. 52 Maximum parsimony: All else being equal, a trait is assumed to only evolve once and be present in all of the descendant organisms. SKILL Create A Tree: Animal Selected Vertebrates Opposable 4-chamber Amniotic Thumb heart lungs Spinal egg colum n Chimpanzee 1 1 1 1 1 Mouse 0 1 1 1 1 Turtle 0 0 1 1 1 Frog 0 0 0 1 1 Fish 0 0 0 0 1 Lamprey 0 0 0 0 0 Continual revision: As more data is gathered, the phylogenetic relationships among organisms are continually revised to incorporate that data. 53 Role of computers: Computer analysis is needed to determine the similarities in DNA/protein sequence information, as the amount of data is beyond the human capacity to analyze quickly. 1.7: Speciation and extinction have occurred throughout the Earth’s history. (EK1.C.1) 1. Speciation Concepts Species concepts: Not discussed by Darwin. “Biological Species”: A group of organisms that are capable of successfully reproducing with eachother. Certain limitations (bacteria? It’s testable! Triceratops?): Have resulted in other definitions of the term. Speciation rates: How quickly do new species evolve? 54 Gradualism: species are the product of slowly accumulating, small evolutionary changes. Punctuated equilibrium: species undergo long periods of very little change, followed by rapid, large evolutionary changes. Evidence for both. Example: Major Extinctions- eventually all species go extinct. There are several periods of greatly increased rates of extinction through the fossil record. Following these Mass extinctions, the species that survive quickly diversify and occupy the vacated niches left open. Adaptive radiation: When one species evolves in to many species that occupy a variety of available niches. Common after Mass extinctions (consider the mammals after the dinosaurs), or when organisms are isolated on islands (consider the Galapagos finches) 1.8: Speciation may occur when two populations become reproductively isolated from each other. (EK1.C.2) 55 1. Speciation process Reproductive Isolation: If a species is a group of interbreeding organisms, than speciation occurs when a group of organisms can no longer breed with any other individuals. Does the speciation occur when the organisms are physically separated from their parent population (allopatric), or does it occur when the organisms are physically in contact with their parent population (sympatric). Species Barriers: Anything that separates one species from another: Pre-zygotic Barriers: Prevent gametes from combining into a fertilized zygote Physical Barriers- geographical Temporal Barriers- time of day/seasonal Behavioral Barriers- mating rituals Mechanical Barriers- incompatibility of reproductive structures Chemical Barriers- incompatibility of proteins on gametes 56 Post-zygotic Barriers: Prevent the hybrid zygote from becoming an organism capable of reproducing Reduced viability- Hybrid is frail Reduced fertility- hybrid is sterile Loss of hybrid characters- hybrid loses hybrid traits over generations. Examples: Mules, Apples, Fruit Fly Food Preference Study. 1.9: Populations of organisms continue to evolve. (EK1.C.3) 1. Ongoing evolution of organisms Ongoing evolution of organisms: Evolution is an ongoing process that can be observed and studied in currently living populations of organisms. Examples: Pesticide resistance: The application of pesticides to crops drives the evolution of pesticide resistance among pest organisms, which requires increased pesticide in subsequent applications. Rock Pocket Mice: Coat coloration evolution has been driven by differences in the rock coloration where the populations live. 57 Different black populations have evolved different mechanisms of melanin production (“convergent evolution”) HW eq. applications: HW Equilibrium can be used to analyze how a population is evolving from one generation to the next. 1.10: There are several hypotheses about the natural origin of life on Earth, each with supporting scientific evidence. (EK1.D.1) 1. Origin of life Nature of science re: Testable hypotheses for life’s origin. Two Major hypotheses. Panspermia: Life has an extraterrestrial origin, arriving on a meteor or similar delivery system Abiogenesis: The origin of living systems from non-living components. A scientific hypothesis because it provides testable predictions. Four major “milestones” for life to develop from non-life: Development of the chemicals that living systems are made of. 58 Encapsulation of those chemicals into isolated systems. Development of an information storage molecule that can be inherited. Reproduction of living systems. RNA world: The hypothesis that RNA preceded DNA. Due to RNA’s “double function” in living systems as an information storage molecule and a catalytic molecule. Evolution of metabolic pathways: The development of the major ways that living systems process matter and energy. Glycolysis is thought to have evolved first, as all modern living systems use it as the first energy production pathway. Photosynthesis evolved later, leading to the oxygenation of the atmosphere. Aerobic cellular respiration evolved last, as it requires atmospheric oxygen to be accomplished. Endosymbiosis: The origin of eukaryotic cells from prokaryotic ancestors. Free- living ancestors of eukaryotic mitochondria and chloroplasts were 59 engulfed by eukaryotic ancestors, and a symbiotic arrangement was established. Supported by a lot of evidence. Evolution of multicellularity: Multicellular organisms are able to specialize the structures and functions of their cells to occupy niches that are unavailable to unicellular organisms. Many unicellular organisms have multicellular life stages, and the systems that multicellular organisms use to coordinate and communicate among their cells are evolved from systems that are present in unicellular organisms. Deep Time Considerations: Approximate evolutionary dates (based on current evidence): Formation of the Earth: 4.5 bya Origin of life: Origin of Photosynthesis: ~4 3 Origin of Eukaryotes: Multicellular life: Origin of Animals: bya bya 2 bya 1 bya 600 mya 60 1.11: Scientific evidence from many different disciplines supports models of the origin of life. (EK1.D.2) Discussion 1: Evidence for the Origin of Life Geological evidence (Dating, Banded Iron, etc.): Radioisotope dating allows us to establish the approximate age of fossil evidence. Different events in life’s history have left geological evidence: Banded Iron formations: Deposits of Iron Oxides that coincide with the hypothesized origin of photosynthesis. Fossil Fuels: Massive deposits of fossilized plants and other producers from the Carboniferous period. Miller-Urey Experiments: Starting with simple chemical building blocks, in conditions that were hypothesized to approximate early earth conditions, simple organic molecules were produced (step 1 of abiogenesis). This finding has been replicated in a variety of experimental conditions. Commonalities of all organisms: 61 The commonalities among all living systems support evolutionary descent from a single common ancestor. It is the simplest hypothesis to explain the observation. 62 Domain 2: Matter 2.1: Organisms must exchange matter with the environment to grow, reproduce and maintain organization. (EK2.A.3) 1. Matter exchange The atoms in living systems are common on this planet. Living systems are mostly made of 6 elements and a handful of other “trace” elements. Living systems cycle matter from the environment, turning it in to biologically useful molecules before returning it back to the environment through life processes. Carbon: Carbon is the major structural atom in all organic molecules. The major non-living source of carbon is the atmosphere (as CO2). Carbon is incorporated in to producers through photosynthesis and from there in to living systems through the food chain, and returned back to the environment through cellular respiration, and decomposition. Oxygen: 63 Oxygen is also found in most organic molecules. The major non-living source of oxygen is the atmosphere (as O2) and water (also the major non-living source of hydrogen). Oxygen is incorporated in to cellular respiration and through the food chain. Oxygen is returned back to the environment through photosynthesis. Nitrogen: Nitrogen is found in all proteins and nucleic acids (present in all living systems). The major non-living source of Nitrogen is the atmosphere (as N2). Nitrogen is incorporated in to the food chain through the action of nitrogen-fixing bacteria, which convert N2 in to molecules that can be used by producers (NO3-), and from producers through the food chain. Nitrogen is returned back to the environment through decomposition and the action of denitrifying bacteria. Phosphorous and Sulfur: Phosphorous is found in all nucleic acids. Phosphate groups are used to quickly store and release free energy in cells. found in all proteins. elements is rocks. Sulfur is The major non-living source of these Weathering processes release them in to the 64 soil/water, where they are available for producers to incorporate and then move through the food chain. Decomposition returns them back to the environment. Hydrogen: Hydrogen is a major component of all organic molecules. the most common atom in the Universe. It is It enters biological systems largely bonded to oxygen in water, and is returned to the environment by decomposition and water release. 2. Properties of Water Water is a major component of all living systems. Water molecules occupy most of a cell’s volume. Water has major properties that living systems require. These properties are due to water’s polarity (unequal sharing of electrons, leading to a partially positive and partially negative charge). 65 Hydrogen Bonds: the polarity of water results in attractive forces between the oxygen and hydrogen atoms of neighboring water molecules. This makes water cohesive (attracted to itself) and adhesive (attractive to anything else that has positive or negative charges). Uses: Transpiration pull, dissolving of substances. Temperature buffering: The presence of hydrogen bonds requires more energy to be absorbed or released by a given amount of water to change the temperature of that water. the specific heat of a substance. heat. This is known as Water has a very high specific Water can absorb or release a comparatively large amount of energy before its temperature changes. Water will keep the environment warmer when the air cools (because it has a larger store of energy to release), and keep the environment cooler when the air warms (because it can absorb a larger amount of energy without changing temperature). Uses: Insulating the ocean. Keeping living systems within homeostatic temperature ranges. 3. MATH Skills: pH 66 Water dissociates. A water molecule can be pulled apart into a proton (H+) and a Hydroxide ion (OH-) by other water molecules. This happens ~ once for every 10,000,000 (107) water molecules. The products of the H+ and OH- ion concentrations in an aqueous solution is a physical constant (10-14). As the concentration of one ion increases, the other must decrease. In pure water, the concentrations of H+ and OH- are each 10-7 (one in 10 million). Acids: substances that increase the amount of H+ ions in an aqueous solution. This will cause the concentration of OH- ions to decrease. Bases: Substances that decrease the amount of H+ ions in an aqueous solution. This will cause the concentration of H+ ions to increase. pH: a measurement of the concentration of H+ ions in a solution. pH = - log [H+] This is a complicated way of saying that we can take the exponent of the pH concentration and negate it (make it positive) to state the pH. Acids have a pH lower than 7. Bases have a pH higher than 7. Each whole number on the pH represents a power of 10. A solution with a pH of 5 has a pH that is 100 times more acidic than a solution with a pH of 7. 4. Constraints on Cell Size 67 Living systems are constrained by the need to exchange matter with the environment. Cellular life has an upper and lower limit on the size a cell can be: Lower limit: Constrained by the need to have a certain amount of matter inside the cell in order to keep functioning. Upper limit: Constrained by efficiency. Surface area: Volume ratio- as a three-dimensonal object’s volume increases, so does its surface area. However, volume increases as a cubic function of size, while surface area increases as a squared function. As volume increases, the efficiency of a cell’s material exchange with the environment decreases. Surface Area Increase Adaptations: A variety of structures have evolved to maximize surface area in cells and increase material exchange efficiency 68 Examples: Microvilli: root hairs: Plant root cells Vertebrate intestines 5. MATH Skills: Surface Area:Volume Ratio Being able to calculate the surface area and volume of an object, if provided with its physical dimensions. Use the formula sheet to find relevant equations. Sample problem: Determine the relative efficiency of material exchange for a spherical cell with a radius of 10 μm, and a cubic cell with a side length of 10μm. Determine the surface area:volume ratio Surface Area of cube: 6xarea = 6 x 100um2 = 600um2 Volume of cube: l*w*h = 10 x 10 x 10 = 1000um3 SA:V ratio of cube: 600:1000 = .6 Surface Area of sphere: 4 pi r2 = 4 x 3.14 x 100 = ~1257 um2 Volume of a cube: 4/3 pi r3 = 4/3 * 3.14 * 1000 = ~4189 um3 SA:V ratio of sphere: 1257:4189 = ~.3 Divide the surface area by the volume. Smaller numbers mean more volume per unit of surface area (less efficiency in transport. 69 2.2: The subcomponents of biological molecules and their sequence determine the properties of that molecule. (EK4.A.1) 1. Biological Molecules Living systems are made of four major types of Macromolecule: Carbohydrates- Sugars: function in short-term energy storage and structural support. Lipids- Fats, oils, waxes: function in long-term energy storage, cell membrane structure, and cell signaling. Proteins- function in all cellular processes. Nucleic Acids- DNA and RNA: function in information storage and expression of that information by determining protein sequences. All macromolecules are synthesized via condensation reactions (“dehydration synthesis”- the removal of water) and decomposed via hydrolysis reactions (the addition of water). Except for lipids, Macromolecules are polymers, made of repeating subunits (monomers). Carbohydrates: Monomer: Monosaccharide \ Polymer: polysaccharide 70 Monosaccharides exist as ring structures. Lipids: Steroids, triglycerides, and phospholipids. Proteins: Monomer: Amino acid \ Polymer: polypeptide There are 20 different types of amino acids used in biological systems. They all have the same “Backbone” structure, but differ based on the composition of the “variable group” that is bonded to the central carbon. The bond between two amino acids is referred to as a “peptide” bond. Nucleic Acids: Monomer: nucleotides \ Polymer: nucleic acid There are two types of nucleic acids in biological systems (DNA and RNA). Each one is made of four types of nucleotides, which differ from each other based on the type of nitrogenous base that they contain (DNA: ACTG / RNA: ACUG) 2.3: Variation in molecular units provides cells with a wider range of functions. (EK4.C.1) 71 1. Variation in Biological Molecules- Carbohydrates and Lipids The structure of carbohydrates and lipids determines their function: Ex: Cellulose vs. Amylose (“starch”): Both are polysaccharide polymers of glucose monomers, but they vary in the connections between monomers. As a result, cellulose is not easily digested, while amylose is very easily digested. Ex: Lipid membrane composition: Cells will vary the lipid composition of their membranes in order to respond to changes in temperature. One example- changes in cholesterol concentration to resist increasing and decreasing membrane fluidity. 2. Variation in Biological Molecules- Proteins The structure of proteins determines their function: Proteins are functionally responsible for all cellular processes. They have a wide and diverse variety of functions, which are 72 determined by their structure. Protein structure is discussed at four levels of organization: Primary structure: the sequence of amino acids in a polypeptide. This is determined by DNA. Each amino acid is joined to the next via a peptide bond. Secondary structure: Regular 3D structures that arise in polypeptide chains due to hydrogen bonding between the oxygen, nitrogen, and hydrogen of neighboring amino acids. Since all polypeptides are made of amino acids, these secondary structures are found in all proteins. Tertiary structure: The unique 3D structure (“conformation”) of a particular polypeptide chain, due to the variety of interactions between the variable groups of the amino acids in the polypeptide and the influence of the environment that the polypeptide chain is in. Quarternary structure: Refers to the overall 3D shape of any protein that is composed of more than one polypeptide chain. Not all proteins have this level of structure. 73 Ex: Adult hemoglobin vs. fetal hemoglobin: Differences in the polypeptide chains that comprise fetal hemoglobin give the molecule a higher binding affinity to oxygen, which is necessary since the fetus needs to get its oxygen from its mothers hemoglobin. Ex: Antibody structure: Differences in the “variable” regions of the proteins that comprise antibodies makes it possible to create millions of different possible antibodies. Denaturation: The disruption of the structure of a macromolecule leading to a disruption of its function. Caused by changes to the environment of the molecule. 3. Variation in Biological Molecules- Nucleic Acids Genetic variation leads to molecular variation. DNA sequence determines protein sequence. Protein sequence contributes to protein structure. Protein structure determines 74 protein function. Protein function determines cellular function. Cellular function determines organism function. A change in DNA sequence can affect all levels of function in a living system. Ex: Sickle cell anemia- one nucleotide change leads to one amino acid change. This change alters the function of hemoglobin, and leads to major disease effects in the organism. New functions can arise via genetic duplications. The duplication of genetic information allows for new variations and adaptations. Ex. Antifreeze gene in fish- temperatures. allows fish to live in freezing Evolved from pre-existing genes. 2.4: Cell membranes are selectively permeable due to their structure. (EK2.B.1) 1. Cell Membrane Structure Cell membrane structure and function: 75 The cell membrane is the boundary between the cell and the environment. It also allows the cell to control which substances pass in to and out of the cell. The cell membrane is composed of a phospholipid bilayer with embedded proteins (the “Fluid mosaic” model). Phospholipid structure causes them to have a hydrophilic phosphate group “head”, and hydrophobic fatty acid “tails”. This leads to spontaneous arrangements in water with the phosphate heads pointing toward the water molecules and the fatty acid tails pointing away. One such arrangement is the “bi-layer” seen in cell membranes. The membrane proteins have a variety of functions: transport, cell- cell contact and recognition, anchorage, enzymatic functions, and others. Selective Permeability: Selective permeability is controlled through control of the proteins present in the cell membrane. Only small, non-polar molecules are able to move through the phospholipid bi-layer. All other substances must move through proteins 76 Cell Wall structure and function: Many cells have a cell wall external to the cell membrane (animals and animal-like cells are the only exception). The cell wall is a structure made of structural polysaccharides (cellulose in plants and plant-like cells) that provides structural support, but no dynamic control of transport. 2.5: Growth and dynamic homeostasis are maintained by the constant movement of molecules across membranes. (EK2.B.2) 1. Mechanisms of Cellular Transport Passive Transport: Refers to the diffusion of material, moving from an area of high concentration to an area of low concentration across a membrane. No energy is required by the cell to enable this process. Simple/Facilitated Diffusion: Simple diffusion: movement through the bi-layer. Only small non- polar molecules can do this. 77 Facilitated diffusion: movement through a protein channel. Any molecule that is not small and nonpolar must diffuse through a protein pore in the membrane. Ex: aquaporins. Active Transport: Refers to the movement of material from an area of low concentration to an area of high concentration. This requires energy, in order to work against the natural tendency for molecules to diffuse. Ex: Na/K pump: Cells always use In order to move sodium out of the cell and potassium in to the cell to establish specific concentrations. ATP is used to provide the energy needed to modulate the shape of the pump proteins. Bulk Transport: The movement of large molecules in to or out of the cell. Requires the use of a vesicle. 78 Endocytosis: Exocytosis: Movement of large particles in to the cell. Movement of large particles out of the cell. 2. Analyzing Transport Tonicity: A measurement of the relative concentrations of solute and solvent in two different solutions. Used to be able to determine how a cell (its internal solution) will respond when placed in to different aqueous environments (the external solution). Three types of tonicity relationships: Hypertonic: The solution has more solute (and less solvent) than the one being compared to. Hypotonic: The solution has less solute (and more solvent) than the one being compared to. 79 Isotonic: The solution has the same amount of solute (and solvent) as the one being compared to. Tonicity relationships and effects on cells: Without the expenditure of energy, material will always diffuse from high concentration to low concentration, if it is able to move across the membrane. Solute will always move from hypertonic solutions to hypotonic solutions. Solvent will always move from hypotonic solutions to hypertonic solutions. As long as it is able to. Cells placed in hypertonic solutions will gain solute and lose solvent (water). Plasmolysis: The loss of water Cells placed in hypotonic solutions will lose solute and gain solvent (water). Lysis: The bursting of the cell membrane. Animal-like cells are adapted to exist in isotonic solutions. 80 Plant-like cells are adapted to exist in hypotonic solutions (the presence of the cell wall prevents lysis). 3. MATH Skills: Water Potential: Water Potential and Solute Potential A measurement of how likely water is to diffuse (“osmosis”) in to an area. Water will move from areas of higher water potential to areas of lower water potential. Pure water is assigned a water potential of 0 (no net diffusion) Water potential depends on several different factors, but we only focus on the pressure difference (the “pressure potential”) and the tonicity (the “solute potential”) as they are the only significant factors in biological systems. Water potential equation: 81 Ψ – Water Potential Ψp – pressure potential Ψs – solute potential The typical units of water potential are bars aka torr aka mmHg in a barometer. In any open air system with no active pressure generation, the pressure potential will be zero, and the water potential will depend entirely upon the solute potential. This is the typical case when investigating diffusion and osmosis in the lab. Solute potential calculation i = ionization constant for the solute (1.0 for sucrose, 2.0 for NaCl, etc.) C = molar concentration of the solute R= pressure constant 0.0831 liter bars/mole K T= temperature in Kelvin (C + 273) 82 Sample Problem: Determine which of the following solutions will gain the most water if placed in to a sample of pure water in a piece of dialysis tubing at the temperature indicated. Assume all samples are at atmospheric pressure: Solution: Solute: Tonicity: Temperature A Sucrose 2M 298K B NaCl 1M 290K C Glucose 1M 300K Water Potential Solution A = -1.0 * 2M * .0831 * 298K = ~ -49.5 bars Water Potential Solution B = -2.0 * 1M * .0831 * 290K = ~ - 48.2 bars Water Potential Solution C = -1.0 * 1M * .0831 * 300K = ~ - 24.9 bars Expect C to gain the most water, since water potential is most negative. 2.6: Eukaryotic cells maintain internal membranes that partition the cell into specialized regions. (EK2.B.3) 83 1. Cellular Compartmentalization Compartmentalization: Increased compartmentalization allows for increased control of life processes, and increased efficiency of those processes, as different components of the cell can function in different environmental conditions. Different areas of the cell can be specialized to perform different processes. Eukaryotes vs. Prokaryotes: The major difference between the organization of eukaryotes and prokaryotes is the amount of internal compartmentalization. Prokaryotes: The vast majority of life on earth. internal compartments (“organelles”). Possess no Unicellular and smaller than eukaryotic cells. Eukaryotes: Contain many different internal compartments separated from the rest of the cell by membrane (“organelles”). Most notably, the nucleus, but many others. Two major types: “plant-like” and “animal-like”, according to how they process 84 energy. Most eukaryotes are unicellular, too, but all multicellular organisms (fungi, animals, and plants) are eukaryotes. Plant-like vs. Animal-like differences: Plant-like cells contain chloroplasts (for photosynthesis), a large central vacuole, and an external cell wall. Animals do not have any of these organelles. 2. Major Eukaryotic Organelles Major Eukaryotic Organelles: Nuclear membrane: rest of the cell. Surrounds and separates the DNA from the Contains pores to allow material to enter and exit the nucleus and interact with the DNA. Golgi Apparatus: A series of flattened membranous disks. Receive vesicles of membrane and protein from the endoplasmic reticulum and modify those proteins prior to routing them to their final destinations. Also the site of polysaccharide production for the cell. 85 Endoplasmic Reticulum: throughout the cell. A series of membranous channels that run Responsible for producing all membrane used by the cell and lot of roles in intercellular transport. Divided in to two parts: Rough ER: smooth and rough Named because it is covered in “bound” ribosomes. The ribosomes are producing proteins that will be embedded in membrane or exported from the cell. Smooth ER: Not covered in ribosomes. Involved in toxin detoxification. Mitochondria: The site of aerobic cellular respiration. A double- membrane structure, with a highly-folded inner membrane (a surface area adaptation). Chloroplasts: The site of photosynthesis. A double-membrane structure with a highly-folded inner membrane (a surface area adaptation). 2.7: The structure and function of subcellular components, and their interactions, provide essential cellular processes. (EK4.A.2) 86 1. Organelle structure and function- information processing Ribosome structure and function: All cells contain ribosomes. Ribosomes are made of two subunits of RNA and protein. They are able to assemble and disassemble as required by the cell. In eukaryotes they are able to associate and disassociate with the endoplasmic reticulum. Endomembrane System: The cellular system by which the information in DNA is expressed and incorporated in to cellular processes: Nucleus ER Golgi final destination (membrane or export). 2. Organelle structure and function- matter and energy processing Mitochondria: Mitochondria have a double membrane which allows for separation of different processes that take place in the mitochondria. 87 The highly folded inner membrane (the “cristae”) contains many copies of the enzymes needed to produce ATP by the cell, with maximized surface area. Chloroplasts: Chloroplasts have a double outer membrane with inner membranous stacks called “thylakoids”. The membrane of the thylakoid contains many copies of the enzymes and chlorophyll needed to produce chemical energy from solar radiation during the first part of photosynthesis (the “light reactions”). The inside of the thylakoids (the “stroma”) contain the enzymes needed to produce chemical compounds during the second part of photosynthesis (Carbon fixation). Lysosomes: Lysosomes are membrane-enclosed sacs that contain collections of digestive, hydrolytic enzymes. Lysosomes serve roles in digestion of molecules, recycling a cell’s damaged components, and programmed cell death. 88 Vacuoles: A vacuole is a membrane-bound sac that stores material. Plants have a large central vacuole that increases the cells surface area: volume ratio by decreasing the active volume of the cell. 89 Domain 3: Energy 3.1: All living systems require constant input of free energy. (EK2.A.1) 1. Bioenergetic Theory The First law of thermodynamics: Energy cannot be created or destroyed, only transformed. Consequences for living systems: Living systems need to continually acquire and transform energy in order to do the work necessary to remain alive (“metabolism”). “Free energy”: The energy available in a system to do work. The Second law of thermodynamics: Every time energy is transformed, the entropy (“disorder”) of the universe increases. Consequences for living systems: In order to increase their internal order, or maintain it, living systems must process more ordered forms of matter in to less ordered forms of matter. 90 Living systems are “open” systems: Material and energy move in to living systems from the environment. Following processing, living systems return matter and energy back to the environment in less ordered forms. In order to increase control of energy processing, living systems produce free energy in multiple-step pathways, mediated by enzyme catalysts (which lower the energy required to cause a chemical reaction to occur). Decreases in energy processing by living systems results in disease/death, and an increase in the entropy of the living system. 2. MATH Skills: Gibbs Free Energy Used to determine if a process can occur spontaneously or nonspontaneously. 91 If free energy is released during a process (“exergonic”) it will occur spontaneously (no energy needed to sustain the reaction following activation. If free energy is absorbed during a process (“endergonic”), it will occur non-spontaneously (energy will be continually needed to sustain the reaction. ΔG= change in free energy (- = exergonic, + = endergonic) ΔH= change in enthalpy for the reaction (- = exothermic, + = endothermic) T = kelvin temperature ΔS = change in entropy (+ = entropy increase, - = entropy decrease) To solve problems, you’ll need to be given values. Exothermic reactions that increase entropy are always exergonic (Ex. Cellular Respiration) 92 Endothermic reactions that decrease entropy are always endergonic (Ex. Photosynthesis). Other reactions will be spontaneous or not depending on the temperature at which they occur. Sample Problem: Determine which of the following reactions will occur spontaneously at a temperature of 298K, justify your answer mathematically: Reaction 1: A + B AB Delta H: +245 KJ/mol Delta S: -.02 KJ/K Reaction 2: BC B + C Delta H: -334 KJ/mol Delta S: +.12 KJ/K 93 Reaction 1 Delta G= + 245 – (298*-.02) = + 250.96 Reaction 2 Delta G = -334 – (298*.12) = - 369.76 3. Metabolic Strategies Uses of Biological free energy: To accomplish cellular work (“metabolism”). Living systems use free energy to repair themselves (maintain order), grow (increase their order), and reproduce (transmit order through time). Catabolism: Decomposition vs. Anabolism: Synthesis Metabolic strategies: The strategy an organism uses to acquire and process free energy will have consequences for the life cycle of that organism. Ex.: endothermy & ectothermy Ectotherms: Organisms that conform their internal temperature to the temperature of the environment. Requires that an organism only be metabolically active when the external temperature allows, but benefits the organism by not requiring any energy expenditure 94 to maintain internal temperature. Examples: All organisms except for birds and mammals. Endotherms: Organisms that maintain a metabolically optimal internal temperature regardless of the external temperature. Requires that an organism use free energy to maintain internal temperature, but benefits the organism by allowing a high metabolism, regardless of the external temperature. Examples: birds and mammals. Effects of Body size on metabolic rate: Larger organisms require less energy production per unit of mass than smaller organisms. This is due mainly to the decreased surface area:volume ratio in larger organisms, leading to decreased loss of heat energy to the environment. Free energy considerations and reproduction: Reproductive strategies are optimized for particular free energy considerations. 95 Example: seasonal reproduction- Organisms reproduce during particular times of the year to make sure that they have enough free energy stored to allow for the energy investment involved in reproduction, and to provide the offspring with the free energy necessary to grow following birth. Consequences of insufficient free energy production for individuals, populations, ecosystems: All levels of biological systems are affected by the amount of free energy available. Individuals will suffer disease/death if they do not get sufficient free energy as there is less energy to maintain internal order. Populations will decrease in reproductive output, and decline in numbers as there is less energy being used to maintain the number of individuals through reproduction. Ecosystems will decrease in complexity and productivity, as there is less energy moving through them. 4. MATH Skills: Coefficient Q10 96 t2 = higher temperature t1 = lower temperature k2= metabolic rate at higher temperature k1= metabolic rate at lower temperature Q10 = the factor by which the reaction rate increases when the temperature is raised by ten degrees. Q10 tells us how a particular process will be affected by a 10 degree change in temperature. Most biological processes have a Q10 value between 2 and 3 Sample Problem: Data taken to determine the effect of temperature on the rate of respiration in a goldfish is given in the table below. Calculate the Q10 value for this data. Temperature (°C) Heartbeats per minute 97 20 18 25 32 Solution: Q10 = (32/18)(10/5) = ~1.78 2 = ~3.178 3.2: Interactions between molecules affect their structure and function. (EK4.B.1) 1. Enzyme structure and function The relationship between structure and function: Because the structure of a molecule allows it to accomplish its functions, changes to the structure of a molecule will also affect its function. These changes can have a negative or positive effect on the function of a molecule. Molecules can interact with other molecules or their environment in order to change their structures. 98 Enzymes: Enzymes are reaction catalysts in biological systems. Most enzymes are proteins, though RNA (“ribozymes”) also have some catalytic functions. Catalyst: any substance that increases the rate of a chemical reaction by lowering the activation energy of that reaction while not participating in the reaction. Enzymes work by physically positioning reactants (“substrate”) in orientations that increase the likelihood of chemical bonds being broken or formed. Enzymes are highly specific for the substrates that they interact with. Typically, the name of an enzyme tells you about its substrate in the first part of its name, and ends in –ase. Ex. Protease, lipase, polymerase. The induced fit model of enzyme function: 99 In order to catalyze a reaction, substrate molecules will physically bind to an area of the enzyme called the “active site”. The binding of the substrate to the active site will cause the conformation of the enzyme to change slightly, catalyzing the reaction. co-factors/co-enzymes: In order to function, many enzymes require organic (co-enzymes: “vitamins”) or inorganic (co-factors: “minerals”) groups of atoms to be complexed with the enzyme. Ex. Many enzymes involved in interacting with DNA require zinc 2+ ions as co-factors. 2. Regulation of Enzyme Activity Other molecules can affect enzyme structure and function: Competitive interactions: Refer to any substance that occupies the active site of an enzyme that is not the substrate of that enzyme. These interactions will inhibit enzyme function. Non-Competitive (“allosteric”) Interactions: 100 Refer to any substance that binds to an area of an enzyme that is not the active site and by doing so affects the function of the enzyme (usually by changing the shape of the active site). In this way, the end product of a metabolic pathway can influence the continuation of that pathway by interacting with an enzyme that catalyzes an earlier step in the pathway (“feedback inhibition”) Environmental conditions affect enzyme structure and function: The local environment of the enzyme can affect the shape of the enzyme, which will affect the function of the enzyme. Major environmental factors include temperature, pH, and salt concentration. Other environmental variables can affect the function of the enzyme by increasing or limiting enzyme-substrate binding. Major factors that do this include enzyme concentration, and substrate concentration. Enzyme action can be measured in many different ways: Typical methods include: 101 Appearance of a product. Disappearance of a substrate. Indirect analogs: color change, change in temperature, etc. 3.3: Organisms capture and store free energy for use in biological processes. (EK2.A.2) 1. Energy Processing Autotrophic nutrition: Autotrophs (“self-feeders”) are able to use energy from the environment to convert inorganic molecules in to organic compounds where free energy is stored. They are the producers in all food chains on Earth. Photosynthetic organisms: Use visible light energy to convert water and carbon dioxide in to oxygen gas (waste product) and organic compounds (sugar precursor molecules). Examples: all plants, and phytoplankton (the major producers on the planet). Chemosynthetic organisms: Use high energy inorganic compounds to convert carbon dioxide and water in to organic compounds. The specific high energy compounds depend on the organism. 102 Example: hydrothermal vent producers: Use H2S as their high energy compound, which is released from the vents. Heterotrophic nutrition: Heterotrophs release free energy from organic compounds (from the food chain, either autotrophs or other heterotrophs), and convert those organic compounds in to inorganic compounds. Anaerobic heterotrophs: release free energy. anaerobes: Do not require oxygen in order to Examples: many bacteria, yeast (facultative can do it if they have to). Aerobic heterotrophs: Use oxygen to release free energy. Release ~20X more free energy from food molecules than anaerobes do. Examples: all multicellular fungi and animals. Electron shuttles: Biological energy production utilizes reduction/oxidation reactions. Electrons are taken from some molecules (oxidation) and transferred to other molecules (oxidation). directly. This does not happen The transfer of electrons occurs via “electron shuttle” molecules, which can hold electrons when they are taken from 103 molecules and release them to other molecules when needed. This transfer of electrons also bring protons along for the ride in Biological systems. Examples: NAD+/NADH, FAD/FADH2, NADP+/NADPH ATP in Metabolism: ATP is a short-term free energy storage molecule used in all biological systems. When a cell releases free energy from a food molecule (or incorporates it from sunlight), that free energy is used to turn a molecule of ADP (2 phosphates) in to a molecule of ATP (3 phosphates). The bond between the 2nd and 3rd phosphate is very unstable and easily broken. When it is broken, the free energy that is released is used by biological systems to power cellular work. 2. Photoautotrophic nutrition- light reactions Photosynthesis is a two-part process: The light reactions Carbon Fixation 104 Mechanisms of photosynthesis- light reactions Light is energy: photons of specific wavelengths are used in the light reactions. The light reactions occur in specialized collections of proteins and chlorophyll called photosystems. These photosystems are embedded in the thylakoid membranes of chloroplasts. During the light reactions, light energy is used to remove electrons from chlorophyll, and use those electrons to produce ATP and NADPH. When excited by photons, chlorophyll sends an electron in to the excited state. This excited electron then enters an electron transport chain of proteins. Water supplies photosystem II with replacement electrons, and converts the water in to protons and molecular oxygen. 105 Non-cyclic electron flow produces NADPH: The ETC connects two different photosystems (PS II and PSI). Electrons produced at PSII move to PSI. Electrons produced at PSI either cycle back in to the ETC, and back to PSI, or they are used by an enzyme to produce NADPH from NAD+ Chemiosmosis produces ATP: As Electrons move through the ETC, the free energy that is released is used by the proteins in the chain to actively transport protons from the stroma into the thylakoid space. The high concentration of protons in the thylakoid space provides the free energy needed to produce ATP. The only way protons can diffuse back in to the stroma is through the “motor protein” ATP synthase. 3. Photoautotrophic nutrition 2- Carbon fixation Mechanisms of photosynthesis- Carbon fixation Carbon fixation happens in the stroma of the chloroplasts, controlled by a collection of enzymes that modulate the “Calvin cycle”. 106 During carbon fixation, the ATP and NADPH produced during the light reactions will be used to drive the incorporation of carbon dioxide in to an organic sugar building block called G3P. Rubisco is the enzyme that incorporates CO2 in to the beginning of the Calvin cycle. During the Calvin cycle, the energy in ATP and the electrons in NADPH are used to reduce 3 5-Carbon molecules of RuBP (plus 3 CO2 from rubisco) in to 6 3-Carbon molecules of G3P. of these is the product of the cycle. One The remaining 5 G3P molecules are then converted back in to 3 5-Carbon RuBP molecules to continue the cycle. G3P can be combined to make sugars, and then used in respiration to produce ATP to power cellular work or serve as raw materials (along with other nutrients) to make lipids, amino acids, or nucleotides. 4. Chemoheterotrophic nutrition- anaerobic cellular respiration 107 Mechanisms of Respiration- glycolysis/fermentation Glycolysis occurs in the cytoplasm, controlled by a collection of enzymes. During glycolysis, 2 ATP are used to convert one 6-Carbon glucose in to 2 3-carbon pyruvate molecules. This process also produces 4 ATP molecules (2 net), and 2 molecules of NADH. Following glycolysis, cells either commit to aerobic cellular respiration, or use a fermentation pathway to oxidize the NADH produced in glycolysis back to NAD+ in order to continue glycolysis. Fermentation will oxidize NADH by reducing pyruvate in to another organic molecule (which depends upon the organism undergoing fermentation). Ex: yeast cells- convert pyruvate in to 1 carbon dioxide molecule and 1 2-Carbon ethanol molecule. Human muscle cells convert pyruvate in to lactic acid. All fermentation end products are more toxic than pyruvate, but the conversion is necessary in order to supply glycolysis with continual NAD+ 108 5. Chemoheterotrophic nutrition: respiration Aerobic cellular Mechanisms of Respiration- Aerobic Cellular Respiration In eukaryotes, ACR occurs in the mitochondrion. Pyruvate must be transported in to the matrix of the mitochondria. It is also converted in to a 2-Carbon acetyl group, which is combined with a molecule of co-enzyme A. Citric Acid Cycle. Acetyl Co-A is the input for the This conversion produces another molecule of NADH The Citric Acid Cycle: The citric acid cycle occurs in the mitochondrial matrix, controlled by a collection of enzymes. During the citric acid cycle, the remaining carbons from glucose (on the acetyl group) are converted in to carbon dioxide (oxidation). This process produces 3 NADH and 1 FADH2, along with 1 ATP per acetyl group. Chemiosmosis produces ATP: 109 Following the citric acid cycle, the NADH and FADH2 that have been produced during all prior parts of cellular respiration are oxidized at electron transport chains embedded in the mitochondrial inner membrane. As Electrons move through the ETC, the free energy that is released is used by the proteins in the chain to actively transport protons from the matrix into the intermembrane space. The high concentration of protons in the intermembrane space provides the free energy needed to produce ATP. The only way protons can diffuse back in to the matrix is through the “motor protein” ATP synthase. Approximately 3 ATP are produced per NADH oxidized. Approximately 2 ATP are produced per FADH2 oxidized. These numbers vary depending on the particular cellular system. At the end of the ETC, the electrons combine with oxygen and protons to produce water. 110 All macromolecules are able to enter cellular respiration pathways, either by being converted in to glucose, or by being converted in to pathway intermediaries. 3.4: Cooperative interactions within organisms promote efficiency in the use of energy and matter. (EK4.B.2) 1. Organizational efficiency in energy processing. Compartmentalization allows for increased efficiency. Cellular compartmentalization: By having different metabolic processes occurring in different cellular compartments, the conditions that those processes occur under can be varied without interfering with other processes. Compartmentalization also allows cells to establish specific locations for specific processes. Example: Mitochondrial/Chloroplast compartmentalization allows for the varied pH/proton sequestration that chemiosmosis will generate. 111 Example: all of the proteins for required metabolic processes are isolated and grouped in to different cellular compartments. This allows for more efficient progression through a metabolic sequence, and also increases the likelihood that any feedback inhibition will be efficient. Ex. Gram-negative bacteria have adapted their cell membranes to compartmentalize processes needed to carry out aerobic cellular respiration and photosynthesis Multicellular compartmentalization: Different systems in a multicellular organism serve different, cooperative purposes for matter and energy processing: Digestive system: Conversion/absorption of complex food molecules in to metabolic inputs (starch in to glucose) Respiratory system: exchange of metabolic gases (oxygen and carbon dioxide) Circulatory system: Delivery of nutrients and removal of waste products from all cells. Excretory system: Removal of waste molecules (water and nitrogenous wastes). 112 All of these systems have different roles in contributing to an overall metabolic goal. Microbial cooperation: Microbial communities diversify in their functions to cooperatively accomplish metabolic tasks. Ex: Animal rumen communities. Ruminants are herbivorous mammals (eg cows) who are able to digest cellulose due to the cooperative actions of microbial communities in their expanded upper digestive tract (the “rumen”) 113 Domain 4: Information 4.1: DNA, and in some cases RNA, is the primary source of heritable information. (EK3.A.1) 1. DNA Structure DNA is the hereditary material in all cells. This understanding is the result of work done by many scientists over the past 100 years. Frederick Griffith: Determined the molecular basis of heredity due to experiments with different strains of bacteria. Avery, McCarty and McLeod: Determined that DNA was the hereditary molecule in cellular systems. Hershey and Chase: Conclusively verified the work of Avery, McCarty, and McLeod. Erwin Chargaff: Determined that the amount of adenine = the amount of thymine, and the amount of guanine = the amount of cytosine in any DNA sample. 114 Watson, Crick, Franklin, and Wilkins: Determined the structure of DNA, and related the structure to the role of DNA in heredity. DNA & RNA Structure and function: DNA and RNA are both nucleic acids made of nucleotide monomers. Nucleotides are comprised of a 5-carbon sugar, a nitrogenous base, and a phosphate group. Nucleotides exist in the cytoplasm as “nucleoside tri-phosphates” (ex ATP, GTP). The extra phosphates provide the energy necessary for a nucleotide to be incorporated in to a nucleic acid. In DNA, the sugar is deoxyribose. In RNA the sugar is ribose. The DNA bases are Adenine, Thymine, Guanine and Cytosine. The RNA bases are Adenine, Uracil, Guanine and Cytosine. RNA is usually single stranded (nucleotides still H-bond with eachother due to bending of the RNA molecule), DNA is usually double stranded. 115 Nucleotides in one strand are covalently bonded between the sugar and phosphate. Nitrogenous bases hydrogen bond with nitrogenous bases on opposite strands. Adenine and thymine/uracil form 2 Hydrogen bonds, Guanine and cytosine form 3 hydrogen bonds. A specific purine (2 rings) always H- Bonds with a specific pyrimidine (1 ring). One strand of a nucleic acid has a directionality, referred to as 5’ and 3’, depending on which carbon of the sugar is exposed at the terminal nucleotide. The two DNA strands are anti-parallel in orientation. The organization of nucleic acids differs in prokaryotes, eukaryotes, and viruses: Prokaryotes: Have one circular chromosome, often with at least one small extra-chromosomal circular plasmid. Eukaryotes: Have many linear chromosomes. Plasmids are rare. 116 Viruses: Have a much smaller amount of genetic material than cells. The viral genome can be DNA or RNA, and single stranded or double stranded. 2. DNA Replication DNA is the information storage molecule in living systems. The “Central Dogma” of molecular biology: Expresses the major flow of information in biological systems. DNA molecules are duplicated (“replication”) to convey information between generations of cells. DNA information is transcribed in to RNA and then translated in to proteins in order to participate in cellular activities. There are other, non-typical paths for information flow seen in viral systems. DNA Replication allows for heritability: 117 DNA is replicated in a semiconservative process. One strand of each molecule serves as the template for the synthesis of the other strand. Each molecule of DNA is composed of one newly synthesized strand and one pre-existing strand. Replication is accomplished through the action of a collection of enzymes that all function together (the “replisome”). Helicase: Opens the DNA helix at the origin sequence for replication. Topoisomerase: Rotates the DNA strand to reduce torsional stress during replication. DNA Polymerase: Responsible for incorporating free nucleotides in to a new strand of DNA. end of a molecule. Can only add nucleotides to the 3’ Makes mistakes. Nucleic acids can only be synthesized in a 5’ -> 3’ direction, which requires one new strand to be made continuously, while the other strand must be synthesized in pieces (“discontinuously”) called “Okazaki fragments”. 118 Ligase: Enzyme that forms covalent bonds between adjacent nucleotides in one strand of DNA. Needed to join the okazaki fragments together. 3. Protein Synthesis Transcription converts DNA sequence information in to RNA sequence Information: RNA polymerase: Enzyme that is responsible for making a strand of RNA (in the 5’ to 3’ direction) from a sequence of DNA. RNA plays several roles in the expression of genetic information: mRNA: RNA sequence of a DNA segment that specifies a polypeptides amino acid sequence. tRNA: Molecules that bring specific amino acids to the ribosome, as dictated by the mRNA sequence. rRNA: Structural components of ribosome subunits. Regulatory RNA: Control gene expression by blocking transcription (RNAi). 119 Translation converts mRNA sequence information in to polypeptide sequences: Translation occurs at the ribosome. The genetic code is interpreted as a series of 3-nucleotide codons (three adjacent nucleotides). There are 64 possible codons, which code for all 20 amino acids, along with a START codon and 3 STOP codons. The Genetic code is redundant, unambiguous, and punctuated. The mRNA interacts with the ribosome to begin translation at the START codon (AUG) closest to the 5’ end of the mRNA. Following initiation, subsequent amino acids are brought to the ribosome as specified by subsequent, adjacent codons. amino acid is transferred to a growing polypeptide chain. Each This is referred to as “elongation” This process continues until the first STOP codon is reached, which triggers “termination”; the release of the polypeptide from the ribosome, and the disassembly of the ribosome. 120 4. Genetic Information Processing & Expression There are differences in gene processing among viruses, prokaryotes, and eukaryotes. Viral gene processing: Some viruses (“retroviruses”) utilize an enzyme called “reverse transcriptase” to convert an RNA genome in to a DNA sequence. This DNA sequence then integrates in to the host cell’s genome to allow for continuing expression of viral components. Prokaryotic gene processing: The lack of a nucleus allows prokaryotes to directly couple transcription and translation. Ribosomes can begin translating an mRNA transcript before RNA polymerase finishes transcribing it from the DNA. Eukaryotic gene processing: Eukaryotes extensively process mRNA transcripts following their production. There are three major post-transcriptional modifications: 121 Addition of a poly-A tail to the 3’ end of the transcript: Aids in transport of the mRNA from the nucleus and increases the longevity of the transcript. Addition of a GTP cap to the 5’ end of the transcript: similar in function to the poly A tail. Removal of introns, and splicing of exons: Introns are interspersed sequences of nucleotides that do not code for functional parts of a polypeptide. They must be removed prior to translation if a cell is going to make a functional protein. The remaining, functional segments (the “exons”) are spliced together to form the mature mRNA transcript. Gene expression contributes to phenotypes through the control of protein activity. Example: enzymes. Enzymes are proteins, and the reactions that they control will only occur if those enzymes are present. Example: Fungal auxotrophs: Mutant strains of fungi with defective copies of enzymes that are involved in metabolic pathways. These strains will only grow if they are provided with intermediary 122 nutrients that enter a metabolic pathway after the action of the defective enzyme. Example: Albinos: The presence or absence of the normal pigments in an organism are due to the presence or absence of particular enzymes responsible for pigment production. 5. Genetic Engineering- Techniques Genetic Engineering allows for direct manipulation of genetic material. This differs from selective breeding because it allows for more targeted control of specific genes. Genetic Engineering Techniques Electrophoresis: separation of DNA molecules based on their size. Utilizes an electrical field and a gel matrix (typically agarose). Smaller DNA molecules migrate through the gel faster than larger molecules. Used to isolate specific genes from within larger samples of DNA, or to visualize differences in DNA sequences among a collection of sequences. 123 Transformation: The direct introduction of DNA sequences in to prokaryotic cells. The DNA sequences are delivered on plasmid vectors which allow for their replication and expression of any genes on them by the prokaryotic cell. Used to engineer prokaryotic cells to produce useful proteins (ex. Human growth hormone). The eukaryotic version of transformation is called “transfection”. Similar purpose, different methodologies. Restriction Enzyme Analysis: Restriction enzymes cut DNA at specific sequences, producing restriction fragments. This enables the isolation of different segments of DNA, for introduction in to plasmids, sequencing, or a variety of other studies. Regions of the human genome associated with identity and diseases will produce different length restriction fragments depending on the sequence of DNA present. Analysis of these Restriction Fragment Length Polymorphisms allows for establishing identity and genetic testing for different diseases. Polymerase Chain Reaction: In vitro DNA replication, using a target sequence of DNA, small primers to bracket the target sequence, a heat-resistant polymerase originally isolated from a thermophillic bacterium (Thermis aquaticus), and a thermal cycler, which oscillates between three different temperatures very rapidly. 124 Through PCR, billions of copies of a DNA sequence can be created in a test tube over the span of a few hours. 6. Genetic Engineering- Applications Many products can be created using genetic engineering: Genetically modified foods: genes for desirable characteristics can be inserted in to the genome of food crops (ex. golden rice) Transgenic animals: genes for desirable characteristics can be inserted in to livestock and other animals (ex. Fluorescent fish) Pharmaceuticals: genes for therapeutic proteins can be inserted in to bacteria to produce large quantities of those proteins (ex. Insulin). Cloned animals: Organisms that do not naturally reproduce asexually can be made to through genome manipulation in the laboratory. There are many ethical considerations associated with genetic engineering: Ethical considerations relate to: Questions of ownership Questions of consent 125 Questions of the definition of life, and what should be legally permissible. 4.2: Changes in genotype can result in changes in phenotype. (EK3.C.1) 1. Mutations Mutations are changes to the genome that can affect phenotypes. Mutations can be positive, negative, or neutral in terms of their affect on an organism’s phenotype. They are caused by exposure to certain chemicals (‘mutagens’), high energy radiation, and through DNA replication (in humans the post-error checking rate of mutations is approximately 1 per 1 billion replicated bases). DNA-level mutations: Substitutions: Change of one base to another base. Can affect the structure of a protein by changing one amino acid, or changing the placement of a stop codon. Called a “point mutation” because it affects only one place in the genome. In/dels: insertion or deletion of a DNA base. Can affect the structure of a protein by changing the amino acid sequence of all subsequent amino acids following the in/del, or changing the 126 placement of a stop codon. Called a “frame shift” mutation because it affects the “reading frame” of the ribosome. Chromosomal mutations: Mistakes that result in large-scale rearrangement of chromosome sections: Duplication Deletion Inversion Translocation Cell division mutations: Mistakes that occur during cell division resulting in extra/missing chromosomes (Down’s Syndrome: Syndrome: trisomy 21, Turner’s monosomy X), or entire sets of chromosomes (polyploidy—really only tolerated in plants) Evolution requires variation. Mutations are the ultimate source of all genetic variation. The fitness of a mutant is not just a function of it’s genome. The environment determines how well adapted a mutant is. 127 Ex. Heterozygote advantage and sickle cell anemia. The sickle cell allele is at a higher frequency in areas where malaria is endemic. Heterozygotes for sickle cell are at an advantage over homozygous non-sickle cell individuals in those areas. In the rest of the world, the heterozygote advantage for sickle cell is not present because malaria is not endemic, and there are other, limiting effects on heterozygote physiology. 4.3: Viral replication results in genetic variation, and viral infection can introduce genetic variation into the hosts. (EK3.C.3) 1. Viral Genetics Viruses are obligate intracellular parasites. Viruses are sub-cellular, and as such are generally not considered to be “living”. They are only active when infecting and replicating inside a host cell, and do not have any external metabolism. Viral anatomy consists of a nucleic acid (DNA or RNA) inside of a protein coat. Some eukaryotic viruses also posses an external lipid envelope. The “generalized” viral “life” cycle (many variations on this theme): 128 Infection: the virus is able to transfer its genetic information in to a host cell. Viruses are specific to particular cell types. This is due to the specificity of the receptors they use to attach to host cells. Replication: the viral genome (and viral polymerases if necessary) utilize host cell materials to manufacture viral proteins and replicate the viral genome. Self-assembly: new viral particles are spontaneously assembled from their components. Release: the new viral particles are released in to the environment to infect new cells. Viruses can evolve very rapidly. Viral polymerases have higher error rates than cellular polymerases. This produces more spontaneous mutations per replication than what is seen in cellular systems. Since more variants are produced every generation, there is a higher likelihood of variants being produced who are better adapted for their environment 129 (which could include features like evading the host immune system). Additionally, when different strains of a virus co-infect the same cell, they can recombine in to variants that have genetic information from each strain. This is the major concern with “bird” and “swine” flu. HIV is an interesting viral example: HIV is a type of retrovirus, a class of viruses that contain an RNA genome. When it infects a cell, HIV uses a viral “reverse transcriptase” to make a DNA copy of its RNA genome. The DNA version of the HIV genome is then spliced in to the host cell genome, where it produces new viruses through the host cell’s normal protein synthesis machinery (RNA polymerase and ribosomes). HIV infects one type of immune system cell (the “Helper T-Cell”), which leads to an eventual collapse of the infected person’s immune system, and the development of AIDS. The body is then susceptible to a series of “opportunistic infections”, which are invariably fatal if not treated. 130 Modern treatments for AIDS include a variety of antiviral drugs that interfere with the HIV life cycle. Viruses are effective at transferring genetic information. Viruses serve as “vectors” for delivering genetic information. Usually this is the viruses own genetic information, but sometimes viruses can be packaged with DNA from the host cell genome (“transduction”). An active avenue of research is looking at using customized viruses to deliver functional genes to populations of cells with genetic defects. There is no “cure” for a viral infection The ability of a virus to remain dormant in a host cell’s genome enables a viral infection to remain latent for long periods of time, before signals trigger the re-emergence of the active virus. In mammals, the immune system recognizes and rapidly responds to recurring viral infections, as long as the virus hasn’t mutated to the point that the immune system can no longer recognize it. 131 4.4: In eukaryotes, heritable information is passed to the next generation via processes that include the cell cycle and mitosis or meiosis plus fertilization. (EK3.A.2) 1. Mitosis All cells reproduce through division (binary fission in bacteria) The Cell Cycle describes the major events of the life of a cell. Interphase vs. M-phase: Interphase is ~95% of the cell cycle: G1: The cell is growing and (if it is going to divide), preparing for DNA replication. S: The cell replicates its DNA. G2: The cell repairs replication errors and prepares for division M-phase: G0: Cell division. permanent interphase that non-dividing cells exist in until death. Through the cell cycle, cells are able to serve the life functions of growth (making more cells), repair (replacing damaged cells), and reproduction (making new generations of cells) 132 The behavior of chromosomes during the cell cycle allows for heritability. Mitosis produces two genetically identical “daughter cells”: The events of mitosis are structured to transmit a complete genome to both daughter cells. The stages of mitosis occur as a continual process. Prophase: the replicated chromosomes condense. Chromosomes are tightly wound DNA molecules, wrapped around histone proteins. Packing makes moving easier. Prometaphase: Metaphase: The nuclear envelope breaks down. The chromosomes align at the middle of the cell, attached to the mitotic spindle Anaphase: The chromosomes are separated at the centromere and migrate to opposite poles of the cell. Telophase: the chromosomes decondense, a nuclear envelope reforms around each set. Cytokinesis: the cell membrane is cleaved, producing two cells. 2. Cell Cycle Control 133 Mitosis is under strict cellular control. In order to move through the stages of the cell cycle, a cell needs to pass through a series of “checkpoints” (internal conditions must be appropriate). Cancer: the result of loss of the cell-cycle controls in a cell. Will inevitably result in the death of the organism. Cells are able to stop dividing, start dividing again if given proper signals from other cells, and remain at particular stages of the cell cycle for long periods of time. The control of the cell cycle is accomplished through the action of proteins: Example: Mitosis Promoting Factor (MPF)- controls the “Mitosis checkpoint” of the cell. enter mitosis. proteins. If MPF is not present, a cell will not MPF is the result of the combination of two other One (Cdk- cyclin dependent kinase) is always present in the cell. The other one (cyclin) is only made once a cell has met the criteria needed to enter mitosis. External signals also control the cell cycle: 134 Example: Platelet Derived Growth Factor (PDGF)- Released at the site of tissue injury by platelets. Causes non-dividing cells to begin dividing to repair damaged tissue. 3. Meiosis Meiosis produces cells with half of the normal amount of genetic material. Sexual reproduction requires that two “haploid” cells combine to produce one “diploid” cell during fertilization. Meiosis is the process that produces these haploid “gametes”. The events of meiosis are similar to those of mitosis with a few key differences: There are two sequential rounds of cell division: Cells progress through meiosis 1 (PMAT1) and then through meiosis 2 (PMAT2) without replicating DNA in-between. become two cells. After meiosis 1, one cell has After meiosis 2, each of the two cells produced in meiosis 1 have become two cells. At the end of meiosis, one diploid cell has become four haploid cells. 135 Crossing over: During prophase 1, the homologous pairs of chromosomes physically associate with each other and exchange genetic material. This results in every chromosome being a unique combination of the genetic material from both members of the homologous pair. Independent Assortment: During metaphase 1, chromosomes line up in the middle of the cell still attached to their homologs. The orientation of one pair of chromosomes has no effect on the orientation of any other pair. Meiosis and the sexual life cycle generate tremendous variation. Variation due to independent assortment: expressed as 2^n. In humans = ~8,000,000 possible orientations of our 23 chromosomes. Variation due to fertilization: expressed as (2^n)(2^n). In humans = ~70,000,000,000,000 possible combinations. Variation due to Crossing Over: “functionally infinite” combinations of genetic information. 136 3. Chromosomal Disorders Many Genetic Disorders are explained by mistakes during the cell cycle. Non-disjunction: Due to problems with the separation of chromosomes during meiosis, cells can be produced with extra chromosomes or missing chromosomes. The majority of these errors are fatal to the developing organism, but some are tolerated, leading to diseases. Ex. Down’s Syndrome: chromosome 21. Results from having three copies of Has wide-ranging effects on the physiology of the person. Ex. Klinefelter’s Syndrome: Results from having two X chromosomes and one Y chromosome. Individuals are phenotypically “male” but have some female secondary sexual characteristics, and are sterile. Genetic tests are available, but pose ethical questions. 137 Karyotype analysis: chromosomes. isolation and visualization of an individual’s Can be used to diagnose non-disjunction related genetic conditions. Modern genetic tests can also determine a host of other genetic diseases that are not able to be visualized on a karyotype (Ex. Huntington’s Chorea) This information can be used by parents to determine if they would like to continue a pregnancy, or prepare for a baby with a medical condition. There are major ethical considerations that accompany these decisions. Questions of reproductive rights, and issues of ownership of genetic information, privacy, and historical contexts (e.g. eugenics). 4.5: The chromosomal basis of inheritance provides an understanding of the pattern of passage (transmission) of genes from parent to offspring. (EK3.A.3) 1. Mendelian Genetics 138 The behavior of chromosomes explains the inheritance of traits. Since genes are on chromosomes, the movements of chromosomes during meiosis will determine the inheritance of the genes they contain. The mathematical analysis of heredity was originally developed by Gregor Mendel, who analyzed the inheritance of traits in pea plants. Mendel developed several major concepts to explain his observations, which have since been related to the behavior of chromosomes, and the relationship between genes and phenotypes: Reminder- Genotype vs. Phenotype Genotype: the alleles that an organism has for a particular trait. Homozygous vs. Heterozygous Phenotype: the trait that an organism shows. Sexually reproducing organisms have two alleles for any trait: One allele is inherited from each parent. 139 “Dominant” vs. “recessive” alleles: the disappearance and then reappearance of certain traits is explained by the fact that these traits are “recessive”. When an allele for a recessive trait is present along with an allele for a dominant trait, the dominant trait will be expressed. In order for a recessive trait to be expressed, both alleles have to be for the recessive trait. Segregation of alleles: During meiosis, each gamete will receive one of the organism’s two alleles. Each organism will produce an equal number of gametes with each allele. Independent Assortment: During meiosis, the segregation of one allele has no effect on the segregation of any other allele, as long as the alleles are on separate chromosomes (“unlinked”). Using these concepts, the inheritance of different characteristics can be predicted based on the 140 proportions of offspring who show different phenotypes for those characteristics. 2. MATH SKILLS: Genetics Probabilities The inheritance of unlinked genes are independent events (similar to multiple flips of a coin). The probability of multiple independent events occurring together is equal to the product of the probabilities of each individual event. Using this, we can determine the probability of a particular group of offspring being produced from a particular cross by determining the individual probabilities for each trait and then multiplying them together. This only works for unlinked genes. Sample Problem: 141 In pea plants the gene for wrinkled seed pods (R) is dominant to the gene for smooth pods (r), and the gene for yellow seeds (Y) is dominant to the gene for green seeds (y). A heterozygous wrinkled, yellow pea plant is crossed with a homozygous smooth, green pea plant. What fraction of its offspring will be smooth and yellow? Cross: RrYy x rryy Offspring: rrY_ ½ * ½ = 1/4 Also, given the limited genotypes for organisms, each type of cross produces characteristic genotypic and phenotypic ratios of offspring. Sample Problem: In a cross between two organisms with the following genotypes: AaBBCcddEe x aaBbCcDdEe What is the probability of getting offspring with the genotype aaBbccDdEE? ½ x ½ x ¼ x ½ x ¼ = 1/128 142 4.6: The inheritance pattern of many traits cannot be explained by simple Mendelian genetics. (EK3.A.4) 1. Mendelian Extensions Many traits are not inherited in simple Mendelian ratios. These traits can involve a lot of different aspects of genetics and physiology, but will always affect the ratios of offspring that are shown. Incomplete Dominance/Codominance: The heterozygote has a different phenotype than either homozygote (ex. Snapdragon color, human blood type) Multiple alleles: Traits have more than two alleles (ex. Human ABO blood types) 2. Non-Mendelian Traits Certain modes of inheritance are non-mendelian. Many traits are linked to certain chromosomes. Mendelian ratios only refer to situations where each gene is inherited independently of each other. 143 The inheritance of these traits depends on the inheritance of these chromosomes. Much of this work was done by TH Morgan’s lab at Columbia in the early 1900’s. Sex linkage: Traits are on the X chromosome. Males will show the recessive characteristic much more often than females. (Ex. white eyes, color blindness) Sex-limited traits: Traits that are only expressed in one sex. (ex. milk production, pattern baldness) Linked genes: Genes that are on the same chromosome will be inherited together, except when crossing over results in recombination. Can be identified by their divergence from Mendelian expected ratios of offspring. Linkage analysis: Can be used to investigate the distance between linked genes on the same chromosome. The greater the distance between two genes on a chromosome, the more often crossing over will occur between them (the greater the frequency of recombinants). Some genes are not inherited through the nuclear genome of a cell. 144 Non-nuclear inheritance refers to the inheritance of genes in the chloroplast and mitochondrial genomes. The chloroplasts and mitochondria of cells are randomly assorted when daughter cells and gametes are being produced, so the traits they carry are not inherited via Mendelian mechanisms. Ex. All mammals inherit their mitochondria from their mother’s egg cell (“matrilineal inheritance”). This can be used to track human ancestry and historical migration. 4.7: Biological systems have multiple processes that increase genetic variation. (EK3.C.2) 1. Generation of Variation All lineages of life have modes of generating genetic variation. Mistakes during DNA replication: Error proofing of DNA is not 100% accurate (thermodynamically impossible). So all DNA will contain some number of mutations due to replication. 145 Horizontal transfer of genetic material in bacteria: Refers to the transmission of genes between members of the same generation of bacteria. Transformation: the uptake of DNA directly from the environment. Transduction: Conjugation: Transposition: The transfer of DNA via viral infection. The transfer of DNA through cell-cell contact The replication and movement of DNA segments separate from replication of the genome. Sexual reproduction: Meiosis and the random nature of fertilization produce functionally infinite combinations of genetic material in sexual reproducing organisms. Sexual reproduction has evolved independently in all major eukaryotic lineages (protists, fungi, animals, and plants). The unifying characteristic is that haploid cells are produced, which unite during fertilization to form a diploid zygote. 4.8: Gene regulation results in differential gene expression, leading to cell specialization. (EK3.B.1) 1. Gene Regulation 146 Regulatory DNA sequences control transcription rates by interacting with regulatory proteins (“transcription factors”): Promoter: sequences of DNA that precede the start of a transcribed gene. polymerase. Recognized by transcription factors and RNA The presence of transcription factors on the promoter make transcription possible. Enhancers: sequences of DNA further upstream than the promoter. The presence of transcription factors on these regions increase the rate of transcription (“up regulation”) Regulatory RNA elements and regulatory proteins. Produced by regulatory genes, and associate with regulatory DNA sequences to allow for transcription. Regulatory DNA sequences serve as molecular “switches”. When the required transcription factors are present, the switch is “on”, and transcription occurs. When the required factors are not present, the switch is “off”, and no transcription occurs. 147 2. Prokaryotic Gene Regulation Prokaryotes control gene expression almost entirely by controlling transcription. The lack of a nucleus makes this efficient. Operons: Groups of genes (“structural genes”) that all contribute to a specific metabolic processes (e.g. digestion of lactose, synthesis of tryptophan). Transcription of all of the genes is regulated by a single promoter. Repressor protein: a regulatory protein that allows/blocks transcription by physically associating/disassociating with a region of the promoter called the “operator”. The modulation between the active and inactive forms of the repressor protein is accomplished through an allosteric interaction with the molecules that are acted upon by the particular metabolic pathway. 148 The operational “logic” of the operon will depend on how necessary the product of the metabolic pathway it controls is for the cell. Ex. The Lac Operon: Involved in the digestion of lactose. “inducible”: usually “off”, only “on” when lactose is present in the cell. The binding of lactose to the repressor protein causes its conformation to change and it to dissociate from the operator. Ex. The Trp Operon: Involved in the synthesis of tryptophan “repressible”: Usually “on”, only “off” when there is an excess of tryptophan present in the cell. The binding of tryptophan to the repressor protein causes its conformation to change and it to associate with the operator. Upregulation: The binding of other regulatory proteins to the operator can increase the rate of transcription of the structural genes to many times above the baseline level. 149 Ex. The Catabolite Activator Protein: When particular energy- producing molecules are low, the cAMP molecular signal associates with the CAP protein. This CAP/cAMP complex associates with the promoter of operons involved in energy production (e.g. the lac operon) and increases the amount of transcription by directly recruiting RNA polymerase to the promoter. Constitutive genes: Genes that are constantly transcribed and are not under major regulatory control. Ex. Ribosomal RNA genes. 3. Eukaryotic Gene Regulation Eukaryotes control gene expression at steps in the process Pre-Transcriptional controls: Access to genes: winding and unwinding of DNA around histone proteins is controlled by enzymatic acetylation and de-acetylation of “tail” regions of the histones. Different types of cells will have different regions of the genome accessible for transcription. 150 Regulation of transcription: Eukaryotes have a wide variety of transcription factors that control transcription (activators and repressors). Different types of cells will have different collections of transcription factors present, to allow the transcription of certain accessible genes and block transcription of others. Post-Transcriptional/Pre-Translational controls: Post-Transcriptional processing: Poly-Adenylation, 5’ capping, and intron splicing all must occur to produce a functional transcript. “alternate splicing”: combining different exons from the same transcript to produce multiple gene products from one transcript. RNA interference (RNAi): the blocking of transcription by small interfering RNA molecules, which bind to specific transcripts and keep the ribosome from being able to translate them. Post-Translational controls: Targeting of polypeptides: signals on the polypeptide determine if the polypeptide will be made in the cytoplasm, or if the ribosome will associate with the ER. 151 Ubiquinone/Proteosome degradation: Tagging of a protein by a ubiquinone molecule will cause the protein to be degraded by a “proteasome” complex. 4. Gene Regulation Phenotype The control of gene expression leads to the control of phenotypes. Phenotypes are the result of protein interactions within the cell. By controlling the production of proteins, cells influence the manifestation of particular phenotypes. Multicellular organisms control cellular differentiation through differential gene expression. The reason that different types of cells are present in a multicellular organism are because different cell types are expressing different genes, which leads to their differing phenotypes. Even genetically identical organisms can have different phenotypes due to differences in gene expression. 152 Ex. Cloned mammals have noticeable phenotypic differences…so do identical twins. 4.9: A variety of intercellular and intracellular signal transmissions mediate gene expression. (EK3.B.2) 1. Signal Control of Gene Expression Signals that are transmitted between and within cells affect gene expression: Example: yeast mating- two mating types: Alpha and a. Each mating type produces a pheromone that interacts with receptors on the cell membrane of the opposite mating type. When this interaction occurs, the genes that control the development of the mating structure (the “shmoo”) in the yeast are activated, and the two mating types fuse together in a fertilization event. Signals that are transmitted between and within cells affect cell function: Example: Hox Genes: present in all animals. Control the development of specific sections of an animal’s body. The presence of Hox transcription factors activate the genes needed to begin the development of a particular segment of an animal’s body (“morphogenesis” and “differentiation”). 153 4.10: Environmental factors influence the expression of the genotype in an organism. (EK4.C.2) 1. Environmental Effects on Phenotype. Phenotype is a product of genotype AND environment: The genome is not a blueprint. It is more like a recipe. An organism’s environment has a major role in determining the organism’s phenotype. The environment can determine phenotype directly: Examples: Hydrangea flower color and soil pH, sex determination in reptiles, The genome can respond to the environment: Example: Seasonal changes in fur color, height and weight in humans, tanning response. “Plasticity”: The interactions between environment and genome can be very complex. It becomes difficult to draw the nature vs. nurture line in many cases. 154 Domain 5: Regulation 5.1: Timing and coordination of specific events are necessary for the normal development of an organism, and these events are regulated by a variety of mechanisms. (EK2.E.1) 1. Development The development of an organism is coordinated by sequential changes in gene expression. Starting as an undifferentiated cell, a multi-cellular organism possessing many different types of cells is produced. Differentiation requires the expression of cell type-specific proteins. Several major processes accomplish differentiation: Pattern formation: Established by positional cues, usually protein gradients (ex. Bicoid, Hox genes). Once body plan axes are established, morphogenesis can begin. Induction: local signals communicated among populations of cells to control their specific development. 155 Environmental cues: Particular molecules and conditions must be present in the local environment for development to proceed properly (ex. The conditions of the uterus, or the role of temperature and moisture in triggering seed development). Evidence from experiments and nature have informed thinking about development: Developmental mutants: mutations in normal developmental pathways lead to malformations in embryonic development. Genetic transplantation experiments: removing/moving regions of the developing embryo to other areas affect pattern formation. The use of reporter genes help to determine which genes are active when and where during organism development. Silencing of specific genes is also important for normal development. It’s not just what you turn on, it’s also what you turn off. The presence of specific microRNA’s during development will prevent certain genes from being expressed. 156 Programmed cell death is required for normal development: Apoptosis: programmed cell death. The cell breaks itself down in to small “blebs”, which are absorbed by surrounding cells. Required for many different aspects of development. example: One digit formation in vertebrates. 5.2: Interactions between external stimuli and regulated gene expression result in specialization of cells, tissues and organs. (EK4.A.3) 1. Differentiation Differentiation is controlled by internal cues (ex. Hox gene products) and external cues (ex. Temperature, or the presence/absence of specific signaling molecules). These all work to regulate protein expression by turning on/off genetic “switches” through controlling the presence/absence of regulatory genes that allow or block transcription. Differentiation of the gene products present in cells leads to structural and functional divergence in the 157 roles that those cells play in the organism (ex. Neuron vs. blood cell) The environment plays a major role in controlling gene expression in mature cells (ex. Changes in UV radiation leading to melanin production, presence of insulin leading to increased glucose intake). 5.3: Organisms use feedback mechanisms to maintain their internal environments and respond to external environmental changes. (EK2.C.1) 1. Feedback Loops Feedback is a universal characteristic of open systems. Feedback loops are seen at all levels of organization in living systems. Negative feedback loops are regulatory in nature. Negative feedback: any situation where the output of a process decreases the occurrence of that process. 158 Example: lac operon- the presence of lactose activates a metabolic pathway that results in the digestion of lactose. Example: Temperature regulation in animals- if animals get too cold, they shiver and increase metabolic rate to generate heat. If animals get too hot, they sweat to cool down. Example: Population growth- As a population grows, resource are depleted to the point where the population can no longer continue to grow. Negative feedback maintains homeostasis. Positive feedback loops are amplifying in nature Positive feedback: any situation where the output of a process increases the occurrence of that process. Example: labor- contractions of the uterus increase pressure on the cervix, which triggers the release of oxytocin by the pituitary gland, which increases the rate of contractions of the uterus. 159 Example: fruit ripening- controlled by the hormone ethylene. As a fruit ripens, it produces more ethylene, which causes the fruit to ripen more rapidly Positive feedback causes transformation in the system. 5.4: Homeostatic mechanisms reflect both common ancestry and divergence due to adaptation in different environments. (EK2.D.2) Discussion 1: Divergence in homeostatic mechanisms is due to the requirements of different environments Examples: Respiratory systems of aquatic and terrestrial animals- external vs. internal respiratory surfaces. Respiration (the exchange of gases) must occur over a thin, moist membrane. Animals that live in aquatic environments can use external structures to respire (gills in fish, or skin in frogs). 160 Animals that live on land must use internal structures to respire (lungs/alveoli in terrestrial tetrapods, spiracles in insects) Nitrogenous waste production in aquatic and terrestrial animalsThe waste products from digesting proteins/nucleic acids must be excreted from the body of the organism. Animals that live in water convert nitrogenous waste directly in to ammonia, which is highly toxic but can be heavily diluted and easily excreted in to the watery environment along. Animals that live on land can not afford to lose as much water when excreting nitrogenous waste. Mammals are able to convert nitrogenous waste into urea which is less toxic than ammonia, so it does not need to be as diluted with water, and can be stored for periodic elimination from the body. Birds, reptiles, and many insects convert nitrogenous waste to uric acid, which is the least toxic, and requires the least dilution with water. 161 Discussion 2: Similarities in homeostatic control systems are due to common ancestry and convergent evolution. Example: Excretory systems in animals. Differences are due All excretory systems in animals involve filtration, reabsorption, secretion, and elimination Examples: Example: Flame bulb/nephridium/kidney(nephron) Circulatory systems in vertebrates- All vertebrate circulatory systems are closed, and use a single heart to pump blood from the body to the respiratory surface and back. Examples: Fish/Reptile/Mammal/Bird- Mammal and Bird 4 chamber heart is an example of convergent evolution. 5.5: Biological systems are affected by disruptions to their dynamic homeostasis. (EK2.D.3) 1. Effects of Disruptions The effects of disruptions to biological systems can be seen at all levels of organization. 162 Disruptions to molecular pathways and cellular structure can adversely affect the homeostasis of the organism: Example: Response to toxins Toxins usually interfere with specific metabolic pathways (ex. Cyanide, Carbon Monoxide), or cause major damage to cells (ex. Snake venom cytotoxins, Concentrated acids). This interference can lead to major injury or death. Example: Dehydration The loss of water causes the tonic environment of cells to become less than optimal for continuing cellular work. The changes in concentrations of molecules can lead to organ failure or death. Disruptions to ecosystems can adversely affect the balance of the ecosystem. Example: Invasive species- If an invasive species is able to outcompete a native species, or place a rapid stress on a native species (e.g. predation), the trophic structure of the ecosystem can be degraded (e.g. 163 decrease in the diversity and abundance of native species) or collapse. Example: Disturbance- Natural disturbances include disasters like fires, earthquakes, etc. These massive, rapid changes to the environment can eliminate major populations from the ecosystem, leading to degradation or collapse of the ecosystem. In all cases, biological systems are able to adjust to the disruptions and rebound (your immune system might be able to produce venom antibodies—Bill Haast, you can drink more water, the populations in an ecosystem can adapt to the presence of an invasive, or an ecosystem can gradually undergo successional changes to return to the pre-disturbance state), IF the disruptions are not too large and too rapid for the homeostatic feedback loops to function. If the disruptions are too large scale, and too rapid, disease, degradation, and death are unavoidable. 5.6: Plants and animals have a variety of chemical defenses against infections that affect dynamic homeostasis. (EK2.D.4) 164 1. Immune Systems. Nonspecific immune responses are found in all multicellular organisms. Plants: Have the ability to recognize pathogens by their effects, and trigger responses that destroy infected tissue and protect uninfected tissue. Invertebrates: Toll-receptor systems that recognize specific molecules usually associated with pathogens (ex. Lipid Acomponent of bacterial cell walls) and trigger immune responses. Vertebrates: External: Internal: Multiple external and internal nonspecific defenses skin, mucous, tears, sweat (lysozyme) Inflammatory response: When pathogens are present inside the body, nearby cells recruit non-specific phagocytic white blood cells to the area of infection. These phagocytes engulf and destroy the pathogens, and present molecules from them (“antigens”) to the specific immune system. 165 Mammals have a highly developed specific immune response: There are two major divisions of the specific immune system. They both involve the lymphocytes (B-cells and T-cells): Cell-mediated response: specific T-cells are developed to target specific antigens on the surface of specific pathogens. These T- cells trigger the destruction of those pathogens, and infected cells through cell-cell interactions. Humoral response: specific B-cells are developed that produce and secrete antibodies that bind to specific antigens on the surface of specific pathogens. These antibodies bind to the antigens on pathogens and target them for destruction by the non-specific immune system, or prevent them from continuing their life cycle. Clonal selection: The process by which specific lymphocytes are produced by the immune system. When presented with a particular antigen, a large variety of lymphocytes are produced, each with a different variation of receptor. Only the cells who react with the specific antigen are allowed to reproduce (the other 166 cells undergo apoptosis). The next generation of lymphocytes undergoes another round. This process continues until a population of lymphocytes with a very specific receptor for a particular antigen is produced. Immunological Memory: Once a specific immune response is generated, a small population of lymphocytes that react to the triggering antigen remain in the lymphatic system for the remainder of the life of the individual. When the same antigen is presented in a subsequent infection, the immune response is much faster and larger, which controls the infection more rapidly. This assumes that the pathogen has not evolved so that its antigens are no longer recognized by the immune system. Memory is also the basis of vaccination, which presents antigens to the immune system without the pathogen present, so that the immune system can develop its response prior to infection. 5.7: Timing and coordination of behavior are regulated by various mechanisms and are important in natural selection. (EK2.E.2) Discussion 1: Changes in the environment are able to cause changes in behavior. 167 The interaction between the environment and internal signals regulate plant responses. Example: Phototropism- Growth in response to light. Controlled by the production and unequal distribution of the hormone auxin. Auxin production is highest in dark cells, which causes them to lengthen. Unequal lengthening results in movement toward light sources. Example: Photoperiodism- Flowering in response to periods of light and dark of specific length. Controlled by phytochrome molecules, which alternate between active and inactive forms as a result of exposure to light for periods of “critical” length. The interaction between the environment and internal signals regulate and synchronize animal responses with the cycles of the environment. Example: circadian rhythms- internal physiological cycles present in all eukaryotes, which last for 24 hours (ex. Sleep/wake cycle). These cycles are present even when environmental cues are removed, but in their absence the cycles lose their calibration to the day/night cycle of the environment (ex. Jet lag). 168 Example: Hibernation/estivation/migration- periods of dormancy, or relocation in response to seasonal variations in productivity of an environment. The interaction between the environment and internal signals regulate and synchronize fungal, bacterial, and protist responses with the cycles of the environment. Example: Fruiting Body Formation- the development of reproductive structures in fungi, some protists, and some bacteria. Controlled by the sensing of appropriate external conditions, and subsequent gene expression. Example: Quorum Sensing- changes in bacterial behavior once a specific population density is reached. (ex. Bioluminescence in V. fisherii) In all cases, interactions between environmental signals and internal signals are required to evoke a particular behavior. In as much as any behavior can be inherited, it can be adapted by natural selection. 169 Domain 6: Communication 6.1: Cell communication processes share common features that reflect a shared evolutionary history. (EK3.D.1) 1. Introduction to Communication Communication requires the generation, transmission, and reception of a signal. In cellular systems, these signals are generally chemical molecules, but can also include direct detection of environmental conditions (eg. Light, sound, temperature). These pathways are referred to as “signal transduction” pathways. All organisms engage have signal transduction pathways: Signal transduction pathways are very important for the continued life of an organism, and are heavily adapted by natural selection. Unicellular signaling pathways: In unicellular organisms, signaling pathways affect the responses of cells to their environment. 170 example- quorum sensing. Involves the generation of chemical signals in response to environmental variables, and they both operate by changing cell activity via regulating gene expression. Multicellular signaling pathways: In multicellular organisms, signaling pathways affect responses that coordinate multiple populations of cells and support the functioning of the organism. Example: glands. epinephrine- a hormone signal produced by the adrenal Triggers different responses in different tissues. In the liver, the reception of the epinephrine molecule causes the breakdown of glycogen in to glucose and the release of that glucose in to the bloodstream. 6.2: Cells communicate with each other through direct contact with other cells or from a distance via chemical signaling. (EK3.D.2) 1. Types of Cellular Signals Cell communication always involves the production, exchange, and receipt of chemical messages. Cell communication through cell-cell contact: 171 Ex. Immune System: Direct contact is needed for activation of the specific immune response (antigen presentation). Ex. Plasmodesmata: Channels in the cell walls of plant-like cells which allow for direct passage of materials and signaling molecules from cell to cell. Cell communication through local signaling: Messages are produced by cells and diffuse to local cell populations Ex. Neurotransmitters: chemicals released by one neuron are received by another neuron over the synaptic space, resulting in the propagation of the signal. Ex. Plant Immune Response: chemicals produced by infected cells are received by nearby cells and result in activation of defense mechanisms in those cells. Cell communication through distance signaling: 172 Endocrine system: the production of endocrine signaling molecules (“hormones”) by glands are then transported through the circulatory system to all other parts of the body. Ex. Insulin: produced by the pancreas, affects all cells in the body (specific liver effects). Ex. Human growth hormone: produced by the pituitary gland in the brain, affects all cells in the body. Ex. Sex hormones: FSH and LH- produced by the pituitary gland, affect the gonads (testes and ovaries). Estrogen, Testosterone- produced by the gonads, affect all cells of the body. 6.3: Signal transduction pathways link signal reception with cellular response. (EK3.D.3) 1. Signal Transduction Pathways Signal transduction begins with reception. Signal reception is accomplished through receptor proteins. Depending on the chemistry of the ligand, receptor proteins will be located at the cell membrane, or 173 in the cytoplasm. Receptor proteins have a diversity of structures, depending on the signal they receive, but there are some general features: An area of the protein that interacts with the signaling molecule (the “ligand”) An area of the protein that transmits (“transduces”) the signal to another protein. When the ligand interacts with the receptor protein, it causes a conformational change in the receptor, which results in the activation of the transduction pathway in the cell. Example: G-protein linked receptor- a membrane receptor. When the ligand binds to the GPL-receptor, the conformational change causes phosphorylation (activation) of a g-protein, which then phosphorylates the next protein in the response pathway, etc. Example: Ligand-gated ion channels- another membrane receptor. When the ligand binds to the channel, the conformational change causes the channel to open, and ions to move freely in to the cell. This change in ion concentration will then trigger cellular responses by changing the shape of various proteins. 174 Transduction of a chemical signal results in the conversion of signal reception in to cellular response. Transduction is accomplished via activation of a protein through phosphorylation, or a change in intracellular conditions (a change in ion concentration). Signal transduction activates a cascading response, which can result in the amplification of the original signal (one ligand - > exponentially increasing activated proteins). The complexities of cellular responses that result from signal reception are due to the interconnected structure of transduction pathways inside a cell. Most signaling pathways involve the activation of “second messenger” response pathways inside a cell. Second messengers are internal signaling molecules, often activated by multiple external signals: Ex. Cyclic AMP -> when present in a cell, activates various catabolic metabolic pathways. 175 The modification of proteins in a signal transduction pathway is akin to turning them “on” and “off”. This is often accomplished by the addition of phosphates to activate proteins (by “kinase” enzymes) and the removal of phosphates to deactivate proteins (by “phosphatases”). Cellular responses involve changes in gene expression, and the activation of already present, inactive proteins. 6.4: Changes in signal transduction pathways can alter cellular response. (EK3.D.4) 1. Alterations to Signaling Pathways Alterations in signal transduction pathways will affect the functioning of cells, and the homeostasis of the organism. Many diseases result from alterations to signal transduction pathways: Ex. Diabetes: Type 1- failure to produce the insulin hormone. Type 2- failure to activate the insulin response in target cells. 176 Immediate effects- inability of the body to effectively regulate blood-glucose concentration. Long-term effects: vascular system problems, poor wound healing, blindness, neurological degeneration, death. Ex. Neurological disease: Parkinson’s Syndrome- death of neurons in the brain that produce the dopamine neurotransmitter. Results in degeneration of the muscular system. Ex. Cancer- failure of cells to respond to the normal apoptosis pathway that should be triggered when cell cycle mutations accumulate. Results in uncontrolled cell growth. Many drugs work by altering signal transduction pathways: Ex. Antihistamines: block the release of histamine signaling molecules by mast cells. Results in decreased inflammatory response. Ex. Birth Control: Provide hormones that prevent ovulation and normal menstrual cycle progression. 6.5: Individuals can act on information and communicate it to others. (EK3.E.1) 177 1. Communication Between Organisms Organisms are able to acquire information about their environment and exchange that information with others. Stimuli: Anything that triggers a response. Stimuli are external to the organism. The ability of organisms to respond to signals from the environment and other organisms (“behavior”) will lead to greater or lesser reproductive success. Ex. Predator warnings- when the presence of a predator in the environment is detected, members of a population will often signal that presence to other members. Ex. Herbivory responses- The detection of chemicals associated with hebivores by plants results in a variety of defenses including the production of toxic chemicals, and the recruitment of herbivore parasites/predators. 178 Animals have highly developed sensory systems, which can detect and communicate via visual, auditory, tactile, chemical, and electrical signals. Ex. Bee Dances: A tactile signal to other bees that relates the position of food sources to the location of the hive. Ex. Swarming Behavior: The result of positive feedback in chemical signaling pathways (“pheromones”). Ex. Prey detection by electrical signals in snakes and fish. As long as behaviors have a genetic component, and increase/decrease survival and reproductive success of organisms, natural selection can adapt them for an organism’s particular environment. Natural selection will favor any behavior that increases survival and reproductive success. Ex. Courtship and mating rituals. Ex. Foraging behaviors in animals. 179 Natural selection will allow for the evolution of cooperative behavior if it increases the fitness of the individual OR genetically related individuals: Ex. Schooling in fish Ex. Colonial insects 6.6: Animals have nervous systems that detect external and internal signals, transmit and integrate information, and produce responses. (EK3.E.2) 1. Neurons The neuron is the structural unit of the nervous system: A neuron is a highly-specialized cell used by the nervous system to detect signals and transmit them to other neurons/response effectors (muscles or glands). Neuron structure allows for neuron function: Dendrites: connect to other neurons or sensory receptors at synapses. Cell Body: Detect signals across the synapse. Contains the majority of the neuron’s organelles. 180 Axon: Conducts an electrochemical signal to the next neuron/effector in the neural pathway. Most axons are surrounded by a layer of highly myelinated Schwann cells that insulate the neuron and increase the rate of signal transmission. Neuron signals move from dendrites to the axon to the nerve terminals. The structure of particular neurons depends on the role of the neuron in the nervous system. Neurons allow for signals to be generated, detected, transmitted, and integrated by animals. Neuron signals are electrochemical “action potentials.” The membrane of a neuron is polarized, with active maintenance of different concentrations of ions inside and outside of the cell (the “resting potential”). cell. Na+ is at a higher concentration outside the K+ is at a higher concentration inside the cell 181 An action potential results from the depolarization of a neuronal membrane’s resting potential. When the membrane is depolarized to a “threshold” potential, voltage gated channels in the axon open, and a rapid exchange of ions occurs: Na+ moves in to the cell, triggering a massive depolarization (inside the cell becomes more positive relative to outside the cell). At the peak of the depolarization, K+ ion channels also open, allowing K+ ions to move out of the cell. Peak depolarization triggers the closing of the Na+ channels, K+ ion channels remain open. As K+ continues to move out of the cell, the membrane becomes hyperpolarized. The action of Na+/K+ pump proteins restores the polarization of the membrane back to the resting potential. Once the resting potential is restored, the neuron can send another action potential. Action potentials are binary (“all or nothing”), self-propagating, and unidirectional. The initial depolarization of the membrane triggers the depolarization of the next area of the membrane. The 182 hyperpolarization following an action potential prevents the action potential from moving backwards along the axon. Myelination greatly increases the speed of action potential transmission, as the signal moves along nodes (“saltatory conduction”). Neurons transmit signals to other neurons across synapses. The arrival of an action potential at the terminal of an axon triggers the release of neurotransmitter molecules in to the synaptic space. Different neurotransmitters will have different effects on different types of neurons. Ex. Acteylcholine: Released by motor neurons at the “neuromuscular junction” (the synapse between them and muscle cells. Ex. Triggers contraction of the muscle Serotonin: Released by neurons in the brain involved in emotional responses. 183 These effects can be excitatory (make the next neuron more likely to send an action potential), or inhibitory (make the next neuron less likely to send an action potential). Transmission of information along neurons will ultimately result in a response (the operation of muscles, or the secretion of signaling molecules by a gland). 2. Nervous Systems Animal Nervous Systems have varying levels of complexity. Evolutionary trends towards centralization and “cephalization” are demonstrated. In vertebrates, the brain is the central unit for integrating nervous system information and coordinating responses. Different regions of the brain serve different functions: Ex. Ex. Medulla/Cerebellum/Cerebrum Right/left hemisphere separation. 184 Ex. Vision and Hearing centers. Ex. Motor movement. Ex. Abstract thought and emotion. Ex. Right/left hemisphere separation. 185 Domain 7: Interactions 7.1: Organisms exhibit complex properties due to interactions between their constituent parts. (EK4.A.4) 1. Physiological Interactions Multicellular organisms are organized into organ systems, which contain organs that work together to accomplish life processes: Ex. Stomach and Small Intestine: Salivary amylase begins the digestion of polysaccharides in to monosaccharides. The stomach combines food with gastric juice, which breaks down food via exposure to hydrochloric acid, and digests proteins through the action of pepsin. of the stomach. Pepsin is only active in the low pH When food is transported to the small intestine the initial section of the small intestine adds pancreatic fluid and bile to the food mixture, the enzymes present in pancreatic fluid continue the digestion of food molecules, and bile emulsifies fat globules. Pancreatic enzymes are only active in the high pH of the small intestine. The later section of the small intestine provides a lengthy surface area over which the end products of digestion are absorbed in to the body. 186 Ex. Plant organs: soil. The roots absorb nutrients and water from the The leaves produce sugars through photosynthesis, the stem allows for exchange of materials between roots and leaves. Organ systems also interact to accomplish life processes: Ex. Respiratory and Circulatory system: The respiratory system allows for oxygen to be absorbed in to the circulatory system. The circulatory system transports oxygen to cells (via hemoglobin in red blood cells) and transports carbon dioxide waste products (as bicarbonate ions) from cells back to the lungs for excretion from the body. If the body is not producing enough energy, the increase in carbon dioxide in the circulatory system causes a drop in pH of the blood which is detected by the nervous system and triggers an increase in respiratory rate. Ex. Nervous and Muscular Systems: The nervous system allows for stimuli to be integrated and a response to be coordinated. The muscular system responds to signals from the nervous system (action potentials), and contracts muscle groups to accomplish motility. 187 Ex. Shoot System & Root System. The vascular bundles provide the leaves with the water necessary to continue photosynthesis via xylem tissue, and provides the nonphotosynthetic cells of the plant with sugars via phloem tissue. The transport of these materials is regulated by the rate of their absorption by other areas of the plant. In leaves, as water evaporates, cohesive attractions move water from the xylem in to the leaf, which “pulls” water through the xylem. As sugar is produced at the leaf (the source), it is transported into phloem. Sugar moves through the phloem until being offloaded at a destination (the sink) 7.2: Organisms respond to changes in their external environments. (EK2.C.2) 1. Responses to the Environment Organisms can respond to changes in their environment by changing behavior: Ex. Fixed action patterns in sticklebacks. Ex. Taxis 188 Organisms can respond to changes in their environment by changing their physiology: Ex. Phototropism Ex. Shivering/Sweating. There is not a clear line between physiology and behavior. Proximate vs. Ultimate explanations: Proximate: Ultimate: How an organism accomplishes a response. Address the evolutionary reasoning for the response. Ex. Imprinting: Proximate Explanation: Immediately after birth, the organism has a “critical period” wherein it will imprint on any moving object near it. Ultimate Explanation: Organisms who imprint are more likely to survive, since the object they are most likely to see is a parent. 7.3: Interactions among living systems and with their environment result in the movement of matter and energy. (EK4.A.6) 189 1. Energy and Matter Acquisition Organisms are adapted to acquire and use matter and energy in their particular environments. These adaptations can be physiological (eg. Gills) or behavioral (eg. Hunting behavior). A well adapted organism for a particular environment may be poorly adapted for another environment (the polar bear in the desert). Energy flows through an ecosystem, matter cycles. Energy is incorporated in to a community by the producers in that community. Producers will usually occupy the greatest biomass in the ecosystem. Primary productivity: the total amount of energy converted into biologically useful forms by producers. GPP (total) vs. NPP (total available to consumers= GPP- metabolism and lost energy). Not all ecosystems are equally productive. The highest productivity is found in locations where there is the most direct sunlight (tropics) and enough nutrients (temperate and cooler bodies of water). Productivity also fluctuates seasonally, and with changes to climate. 190 Within a food chain, only ~10% of energy at any trophic level will be passed on to the next trophic level. Trophic interactions are represented as food chains and foodwebs. Trophic structure can also be represented as pyramids (of biomass, numbers, etc.) Matter cycles between living and non-living components of an ecosystem. All matter cycles involve the movement of matter between abiotic and biotic reservoirs in an ecosystem. Producers are the major pathway for matter to move from the environment to the community. Decomposition is the major way for matter to move from the community back to the environment. Within a community, matter moves through the food chain. Community interactions influence the movement of matter and energy. These influences can be modeled. 191 Competition for resources limits the growth a population. As competition increases, population growth approaches zero (the “logistic model”) The effects of many limiting factors increases as the population density increases (“density-dependent interactions”). Ex. Space, predation, accumulation of waste, etc. Human activity can impact ecosystems locally, regionally, and globally: The exponential growth of the human population has lead to increasing resource consumption in all ecosystems in which the human population lives. Humans now have a major impact on all ecosystems on the planet. This impact is largely detrimental to other organisms in these ecosystems (though not wholly). Human activities have reduced the population sizes of many other organisms through direct consumption and habitat destruction. Many species have gone extinct in part, or in whole due to human impact on the ecosystems in which they live. 192 2: MATH SKILLS: Productivity Calculations It’s mostly accounting. Total energy and matter in to a system has to equal total energy and matter out of the system. Sample problem: A caterpillar consumes 100 kilocalories of energy. It uses 35 kilocalories for cellular respiration, and loses 50 kilocalories as waste (heat and in waste products). Determine the trophic efficiency for its creation of new biomass. Total Energy: 100 Kcal Lost and Respired: 35 + 50 = 85 kcal Total for growth: 15 kcal Efficiency: part/total = 15/100 = .15 (as decimal, 15 as percentage, 3/20 as a fraction, etc.) 193 The formula sheet also provides two conversion factors to use the amount of oxygen gas produced in photosynthesis as an indicator of gross primary productivity: mg O2/L x 0.698 = mL O2 /L mL O2/L x 0.536 = mg carbon fixed/L You are more likely to need to understand that ecosystems with greater productivity are more stable and diverse than ecosystems with less productivity than you will need to do any particularly complicated math in this area of the course. 7.4: All biological systems from cells and organisms to populations, communities and ecosystems are affected by complex biotic and abiotic interactions involving exchange of matter and free energy. (EK2.D.1) 1. Limiting Factors Biotic and abiotic interactions affect all levels of biological systems. 194 Cellular activities are affected by abiotic and biotic interactions: Ex. Cellular proliferation is controlled by the density of cells in the environment, and the amount of resources available. Ex. Biofilms- complex microbial communities that are established and maintained by the actions of all species present in the biofilm. Organism activities are affected by abiotic and biotic interactions: Ex. Predator-prey relationships. The cyclical nature of population changes demonstrates their interdependence. Ex. Resource availability in the environment determines an organism’s behavior (foraging, hibernation, etc.) The stability of populations, communities, and ecosystems are affected by abiotic and biotic interactions: Ex. Resource availability and productivity influence the complexity and diversity of the community, and the size of the populations that comprise the community. 195 Ex. Algal Blooms- Sudden changes in nutrient availability in aquatic environments can lead to a rapid increase in producers, and can trigger the death of other populations in the ecosystem (due to loss of nutrients, or toxins produced by the algae). In all cases: Biotic and abiotic interactions both play roles in affecting biological systems at all levels of organization. These affects can be beneficial, detrimental, or variable in their effects on the system and its state at the time of the interaction. 7.5: The level of variation in a population affects population dynamics. (EK4.C.3) 1. Population Diversity The ability of a population to respond to changes in its environment (“resilience”) is directly related to its genetic diversity. Populations with the least genetic diversity are most at risk for extinction in an ecosystem. Ex. Potato Blight- a major cause of the Great Famine in Ireland. Irish potatoes were all the same 196 strain, greatly decreasing their genetic diversity. all susceptible to the blight fungus. This made them Eventually, resistant strains were developed by incorporating genetic diversity from South American strains. Genetic diversity leads to a diversity of responses among individuals in a population to the same environmental changes. This diversity can be physiological or behavioral. Ex. Black Plague- Already present variations in the gene for the receptor used to infect cells lead to the increased survival of resistant individuals. Ex. Not all animals in a herd stampede when exposed to a panic stimulus. HW eq. can be used to model allele variation in a population, and inform hypotheses about how environmental changes might affect populations. Genetic variation can also be estimated through direct population sampling. Other modes of estimating diversity include phenotype sampling, and fossil record analysis. 7.6: Interactions between and within populations influence patterns of species distribution and abundance. (EK4.B.3) 197 1. Community Interactions Interactions between populations affects the distribution and abundance of organisms. Niche: the total interactions of an organism with its environment. Competition, and predation can limit the distribution and abundance of a population. Competitive exclusion principle: When two species have overlapping requirements in the same ecosystem, one species will outcompete the other for those overlapping resources. Symbiosis can limit or expand the distribution of a population: Mutualism and commensalism are expansive. Parasitism is limiting. A population has properties unique to its level of organization. These properties emerge from the interactions among the individuals who comprise the population with each other and the ecosystem. 198 The distribution and abundance of a population can be disrupted by a variety of environmental changes: Ex. Catastrophes (mount st. Helens). Changes in resource availability (algal blooms, role of iron in aquatic ecosystems). Human activities (introduction of invasive species). Interactions among populations can be modeled, and modeling can inform predictions about the effects of those interactions. It is not possible to accurately model the entirety of interactions between populations and their environment. This makes prediction about the effects of changes on a population inherently uncertain (Hardin’s Law). 7.7: Communities are composed of populations of organisms that interact in complex ways. (EK4.A.5) 1. Measuring Communities Community structure can be measured in different ways: Species composition: the number, distribution, and abundance of the species in the community. 199 Species diversity: the variation within the species and populations in the community. Mathematical models can be used to investigate population growth patterns: Exponential model: Logistic equation: assumes unlimited resources. accounts for the effect of carrying capacity on population growth. Demographic Representations: Age structure of populations Mathematical models can be used to illustrate and investigate population interactions: 2. MATH SKILLS: Population Growth Equations Four Given Equations: Rate of change over time: Population Growth Rate 200 Exponential Model: Logistic Model Using the right equation is half of the battle. Sample Problem: Over the span of one year, there are 20 deaths and 35 births in a population of 200 African elephants. Determine the maximum per capita growth rate of the population. If the carrying capacity for the population in the environment is 300 elephants, determine the number of elephants that can be predicted in the population at the end of the next year, if the intrinsic per capita growth rate does not change. Solution: 201 What is the maximum per capita growth rate for the population? dN/dt = B – D dN/dt = 35 -20 dN/dt = rmaxN 15 = rmax 200 dN/dt = 15 elephants per year. rmax = 15/200 rmax = .075 What will next year’s population be if the carrying capacity is 300 elephants? dN/dt = rmax N (K-N/K) = .075 215 (85/300) = ~4.6 elephants = 4 elephants + 215 = 219 elephants 7.8: The diversity of species within an ecosystem may influence the stability of the ecosystem. (EK4.C.4) 1. Ecosystem Stability Different ecosystems have differing amounts of biodiversity. The more biodiversity present in an ecosystem, the more stable the ecosystem will be, and the more resilient it will be when exposed to changes. 3 levels of biodiversity: The genetic diversity in a population. 202 The number of populations in a community. The diversity of ecosystems in a Biome. Ecosystems are interaction networks (they depend on interactions among the components). possible interactions. The more components, the more The less important any one interaction is to the functioning of the ecosystem. Decreasing biodiversity decreases the number of components in the ecosystem, and increases the reliance on any remaining interactions to keep the ecosystem functional. If something then happens to the relied-upon species, the ecosystem may collapse. Keystone species- species that have a large role in maintaining the structure of the ecosystem. The removal of keystone species from an ecosystem will often result in the collapse of the ecosystem. Ex. Sea otters in kelp forests, sea stars in intertidal systems. 203 Facilitation: When one species makes it possible for other species to occupy an area. Ex. Beavers Producers are required for the continuing functioning of an ecosystem. Changes in the amount of producers affects the total amount of matter and energy moving through the community, which will affect the structure of the community. Limiting Factors: The abiotic and biotic factors present in the environment that are most scarce. Limiting factors limit the growth of populations and the complexity of the environment. Changes in the availability of limiting factors leads to changes in the structure of the community. 7.9: Distribution of local and global ecosystems changes over time. (EK4.B.4) 1. Ecosystem Changes Geological and meteorological events can have large effects on ecosystems (“disturbances”). The effect a disturbance has on an ecosystem is directly proportional to the rate and scale of the disturbance (a forest fire will have a major effect, but not as major as a meteorite impact. 204 Biogeographical data can illustrate the historical effects of disturbances on the history of life on Earth. Ex. Contental drift, meteor impacts Ecosystems have ways to recover from disturbances (ex. Succession). “Nature Abhors a vacuum”. Human impacts have a disproportionate effect on the rate of change in ecosystems. The effects of human impacts are difficult to predict, and far-ranging across all ecological scales (from populations to ecosystems to the biosphere). Ecosystem services: The processes that are accomplished by ecosystems that enable life on Earth to persist (food sources, nutrient recycling, air and water purification). Human activities disrupt the functioning of these ecosystem services, and are contributing to an accelerated extinction rate. Ex. Climate change (and associated ecosystem degradation related to fossil fuel consumption), overexploitation of resources (slash and burn techniques in the rainforest), transmission of emerging diseases (ex. MERS-CoV). The effect of warfare. 205 Human Impacts are accelerating as the population increases. How will human impact affect the long term viability of the biosphere, and our place in it? 206