Which Type Of Selection Would Favor Steadily Increasing Body Size For Wolves In Colder Climates?

• Journal Listing
• CBE Life Sci Educ
• five.12(three); Fall 2013
• PMC3763015

CBE Life Sci Educ. 2013 Autumn; 12(three): 483–493.

Six Classroom Exercises to Teach Natural Option to Undergraduate Biology Students

Steven T. Kalinowski

*Department of Environmental, Montana State University, Bozeman, MT 59717

Mary J. Leonard

^†Department of Instruction, Montana Country University, Bozeman, MT 59717

Tessa Chiliad. Andrews

*Department of Environmental, Montana Country University, Bozeman, MT 59717

Andrea R. Litt

*Department of Ecology, Montana State University, Bozeman, MT 59717

Nancy Pelaez, Monitoring Editor

Received 2012 Jun 12; Revised 2013 January 25; Accustomed 2013 Jan 26.

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Abstract

Students in introductory biology courses frequently take misconceptions regarding natural selection. In this paper, we describe six activities that biological science instructors can use to teach undergraduate students in introductory biology courses how natural choice causes evolution. These activities brainstorm with a lesson introducing students to natural selection and also include discussions on sexual selection, molecular evolution, evolution of complex traits, and the development of behavior. The set of six topics gives students the opportunity to see how natural selection operates in a variety of contexts. Pre- and postinstruction testing showed students' agreement of natural selection increased substantially later on completing this series of learning activities. Testing throughout this unit of measurement showed steadily increasing student agreement, and surveys indicated students enjoyed the activities.

INTRODUCTION

Evolution is the unifying theory of biology (Dobzhansky, 1973 ). It also may be the well-nigh well-supported scientific theory that is rejected by a large proportion of Americans (e.g., Miller et al., 2006 ). This high-profile controversy will be familiar to any teacher teaching evolution in the United States. What many biology instructors may not realize is the theory of evolution is also conceptually difficult for students to understand. In particular, natural selection, the main cause of development, is a challenging concept for many students. This is ironic, because, as many authors have emphasized (e.g., Coyne, 2009 , p. sixteen), natural selection is not a complicated process.

The main reason students often accept trouble understanding natural selection is that they accept misconceptions regarding what causes populations to change. By "misconception," nosotros mean a commonly held thought that is inconsistent with scientific understanding and is resistant to education (sensu Hammer, 1996 ). The origin and cognitive structures that give rise to student misconceptions are the subject of ongoing enquiry (e.g., Hammer, 1996 ; diSessa, 2006 ; Mason, 2007 ; Abrams and Southerland, 2010 ), but the bodily misconceptions students have are well documented (run into Gregory [2009 ] for an excellent introduction to the large literature on student misconceptions relating to natural option). Virtually misconceptions relating to natural selection are variations of the conventionalities that individuals evolve. Students often believe individuals modify considering they need to, because they want to, because the environment changes them, or because they use or do not utilize specific body parts—and that these changes are passed on to offspring (e.g., Brumby, 1984 ; Bishop and Anderson, 1990 ; Nehm and Schonfeld, 2008 ). These misconceptions are frequently similar to Lamarck's (1809) pre-Darwinian theory of evolution.

Student misconceptions regarding natural pick are remarkably resistant to pedagogy (for reviews, see Tanner and Allen, 2005 ; Sinatra et al., 2008 ; Gregory, 2009 ). Bishop and Anderson (1990) showed that half of the students in an introductory biology course left the grade with misconceptions regarding natural selection—even though the lectures and laboratory exercises in the course were designed specifically to address misconceptions. Nehm and Reilly (2007) plant that 70% of biology majors completing an introductory biology course had at to the lowest degree one misconception regarding natural selection—even though the instructors emphasized development as a theme throughout the course and used active-learning exercises extensively to promote learning. Andrews et al. (2011b) studied the effectiveness of introductory biology courses throughout the Us and found that pre- and postinstructional measures of student understanding of natural pick in many courses were statistically identical.

Carefully designed instruction is necessary to help students supercede misconceptions with scientifically supported conceptions. There is a consensus among science educators that active learning is more successful than traditional lectures in developing pupil understanding. Active learning essentially occurs when an instructor stops lecturing and students work on a question or task designed to help them understand a concept (Andrews et al., 2011b ). A classic example is a remember–pair–share discussion, in which students kickoff individually think virtually a question posed by the teacher, then pair up with other students to talk over the question, and finally share answers in a whole-class discussion. Students can larn twice as much when lectures contain discussions than when instructors simply lecture (due east.g., Hake, 1998 ; Knight and Wood, 2005 ; Haak et al., 2011 ). Yet, such active-learning instruction may non be sufficient for replacing persistent misconceptions unless instructors blueprint learning activities that specifically focus on helping students move beyond misconceptions to scientifically supported ideas (Duit and Treagust, 2003 ; Tater and Mason, 2006 ; Andrews et al., 2011b ). Ongoing research continues to identify the most effective ways to help students recognize and change misconceptions (e.g., Vosniadou, 2008 ). Nonetheless, at that place is a consensus that the following instructional practices are useful. Starting time, instructors must help students get aware of what they believe and how to recognize which of their ideas disharmonize with observations or with previous knowledge (Hewson et al., 1998 ; Bransford et al., 2000 ). 2nd, students are more likely to larn any concept, including ones for which they have misconceptions, when they study multiple examples of the concept at work (Catrambone and Holyoak, 1989 ; Mestre, 2003 ; Marton, 2006 ). Tertiary, helping students construct scientific conceptions involves introducing them to scientists' "ways of seeing" (Scott et al., 1991 ); in this case, seeing situations of natural selection in terms of its genetic ground (Kalinowski et al., 2010 ).

There is a shortage of classroom exercises for teaching natural selection that crave the active participation of students, that have been designed specifically to deal with misconceptions, and that have been shown to be effective in promoting a deep understanding of natural selection. In this investigation, we developed and assessed half-dozen classroom exercises for education natural choice to undergraduate students. The main learning goal for the exercises was for students to thoroughly understand natural pick.

METHODS

Participants and Context

We designed and assessed (with institutional review board permission) six lessons relating to natural selection in an introductory course on ecology and evolution during 2011 and 2012. This class was the third in a 3-semester sequence of introductory biology courses for biology majors at Montana State University. The previous courses in this sequence covered physiology, cell biology, biochemistry, and genetics. Classes met three times a week for a 50-min lecture and once a calendar week for a 3-h lab. Twenty lectures in the course were devoted to studying development, including six focused on natural selection.

The student body inverse during our report. In 2011, 41 students enrolled in the grade: 85% were freshman, 12% sophomores, and 3% seniors. In 2012, 47 students were enrolled in the course: 54% were sophomores, 29% were juniors, and 17% were seniors. The large proportion of freshman in 2011 was due to an advising error, but provided us with an unexpected opportunity to test our lessons on ii different sets of students. In both years, virtually all students reported past raised hands they were preparing for careers in the health sciences.

We did not collect demographic information on race either yr. Even so, the educatee body in the Higher of Letters of Sciences at Montana State University is 90% Caucasian, and there is no reason to suspect enrollment in this course was substantially unlike.

Two instructors taught the half dozen learning activities described hither. In 2011, Southward.T.K. taught the entire course. S.T.K. is a tenured professor, who had eight years of teaching feel in 2011. He had taught this course half-dozen times previously. In 2012, T.M.A. taught the 6 lectures that included the learning activities described in this paper. (South.T.Thou. taught the remainder of the course). At the time she taught these classes, T.M.A. was a fourth-yr PhD student whose primary teaching feel consisted of serving every bit a teaching banana for this course the previous yr. Half of T.M.A.'s PhD research related to undergraduate biology education (Andrews et al., 2011a , 2011b ), and then she likely had more pedagogical cognition than virtually graduate students.

Description of the Vi Learning Exercises

We created half dozen learning exercises (Tabular array 1) to teach students how natural pick works and to help correct common misconceptions regarding its functioning. Our exercises focused on students' ideas, drawing from principles of agile learning and conceptual modify education. Although exercises varied, they generally included the following elements: we began by request a thought-provoking question that asked students to business relationship for a state of affairs in which natural selection was occurring, to evaluate whether it was occurring, or to predict what would happen in the situation. We elicited students' initial ideas in response to the question, either in writing or as a class give-and-take. We provided opportunities for students to talk over their ideas with their peers. We explicitly discussed misconceptions in ways intended to help students evaluate them; for example, describing circumstances for which a misconception could not provide an adequate explanation, facilitating students in identifying pros and cons of an explanation, and having students compare alternate explanations. Finally, nosotros emphasized the genetic basis of natural selection past providing genetic data. The learning exercises, described beneath, were given in six lectures (one per lecture) in the lodge described below. Two of these exercises have been described previously: Kalinowski et al. (2010) described the dog-breeding give-and-take, and Andrews et al. (2011a) described and assessed the man evolution discussion.

Tabular array one.

Focal questions for six classroom exercises for pedagogy natural selection, and the approximate amount of fourth dimension required to hash out each question

Exercise	Focal question	Time required
Canis familiaris breeding	How could you turn a pack of wolves into Chihuahuas? Explicate why your strategy would work.	xxx min
Coat color in Oldfield mice	Where did the gene for white fur come from?	10 min
Human development	Are humans evolving? If and so, which trait is changing? Explain.	50 min
Peacock trains	How would y'all test whether peacocks with large spots on their trains accept "good genes"?	xv min
E. coli antibody resistance	What does natural pick predict about the evolution of antibiotic resistance in Due east. coli?	five min
Lemming suicide	Could suicide be an adaptation in lemmings?	15 min

Introduction to Selection: Domestic dog Breeding.

In the first chapter of On the Origin of Species, Darwin described how plants and animals take been bred to accept desired characteristics and used such "bogus" choice as an illustration to innovate how "natural" selection works in nature. The illustration is still powerful, and nosotros used it to introduce our students to natural selection. Nosotros began this word by telling the course that all dog breeds are descended from wolves and asking the course "If you had a bunch of wolves and wanted a Chihuahua, how would y'all create one?" Students discussed the question in pairs and we and so elicited answers from randomly selected students. A "correct" answer for this question is that Chihuahuas can be bred from wolves by selectively breeding minor wolves with short faces and wiry tan hair for many generations. Some students provided this answer, but many proposed raising wolves in a warm environment "then they volition not demand such heavy fur" and providing them with plenty of nutrient "then the wolves get less ambitious and develop smaller teeth." Such responses reveal the misconceptions that the environment causes individuals to evolve and traits evolve from their use or disuse.

Once we recorded a diversity of answers on the board, and the class could run into the need to reconcile the differences expressed, we asked students to comment on the feasibility of each proposal. This created some confusion. Some students clearly understood why raising wolves in domestic environments will not cause them to become Chihuahua-like. Others did not. We resolved the confusion by making a connection to genetics and reminding the class that wolf pups grow upwards to be adult wolves considering they have wolf genes, not considering they are raised in a forest hunting elk.

We emphasized that the selective-breeding plan will work, because when a breeder preferentially chooses small, tan wolves to breed, he or she is selecting wolves with specific DNA sequences. We showed the form DNA sequences for 1 of the genes that differentiate Chihuahuas from wolves. Torso size in wolves and dogs is influenced by variation at the insulin-like growth factor 1 (IGF1) gene. Chihuahuas (and other miniature breeds) accept an adenine at position 44,228,468 in the dog genome, and wolves accept a guanine (Sutter et al., 2007 ).

After we completed the dog-breeding discussion the instructor gave a short lecture describing the requirements for natural option. We emphasized evolution by natural selection would occur if: 1) there was phenotypic variation in a population; two) this variation was heritable, and 3) this variation influenced the reproductive success of individuals. We repeatedly emphasized these three concepts in all our subsequent lectures, and afterward used them in our assessment of pupil learning.

Source of Variation: Coat Color in Oldfield Mice.

One of the challenges for students studying natural selection is that the genetic basis of evolution is invisible. Therefore, discussing how natural choice affects the frequency of DNA sequences in a population should be useful. Coat colour in Oldfield mice, Peromyscus polionotus, makes a useful case written report for how natural option might piece of work in the wild.

We began our give-and-take of coat color in Oldfield mice by describing the natural history of the species. Oldfield mice live in the southeastern United states and generally have dark fur that provides camouflage from owl predation for mice living in forests. In contrast, a subspecies of Oldfield mice, P. p. leucocephalus, lives on the white sand dunes of Santa Rosa Island on the Gulf Coast of Florida and has nearly white fur. Experiments have shown that this coloration protects mice from owls (Kaufman, 1974 ). The difference in fur colour is largely caused by a single nucleotide change in the melanocortin-1 receptor factor. Mice with a thymine at a specific location in the gene have much lighter fur than mice with a cytosine at that location (Hoekstra et al., 2006 ). The mice on Santa Rosa Island are the just mice in the southeastern United States with this genetic variant.

Afterward we presented the preceding material to the class, we asked our students to consider a plausible history for Oldfield mice on Santa Rosa Island. Specifically, we asked them to presume that Oldfield mice colonized Santa Rosa Island when sea levels were lower during the last ice historic period, and that all of the mice colonizing the island had brown fur. We emphasized that being a brown mouse living on white sand dunes put the mouse in danger. Nosotros then asked our students "How did the population of brown mice become white?" Students idea about this question and wrote an answer on an index carte. Then students discussed their answers in modest groups, after which we solicited answers from the entire form. After collecting answers from several groups, we discussed the importance of mutation in creating new phenotypes, and emphasized that mutations randomly changed Dna sequences—and do not necessarily make the changes that organisms desire or need.

Natural Selection: Human Evolution.

Every bit nosotros have discussed above, many students have persistent misconceptions regarding how natural choice operates. Our human evolution discussion is especially effective at eliciting such misconceptions. Andrews et al. (2011b) described in detail how nosotros conduct this discussion, and nosotros will refer the reader to their paper for a full description of the exercise. A cursory description of the discussion should exist adequate hither.

Nosotros begin the word by asking our students "Are humans even so evolving? If so, what trait is irresolute? Explain why or why non." Students discussed this question in groups and then presented answers to the class. Many of the answers lacked any reference to the principles of natural selection. For instance, students answered "people are getting balder" or "computers are making us smarter." We then discussed the likelihood of such changes occurring, given the requirements for natural selection (variation, heritability, differential reproductive success).

Sexual Option: Peacock Trains.

It should non be besides difficult for students to understand how natural selection tin alter the coat color of populations of mice living on sand dunes, but the origin of other traits is harder to sympathise. For example, Darwin struggled to figure out how natural choice could have created highly ornamental traits, such as the peacock's railroad train, that would seem to reduce an private's power to survive. In 1860, he wrote Asa Gray: "the sight of a plume in a peacock's tail … makes me ill!" Darwin eventually proposed elaborate peacock trains could evolve if peahens preferred mating with peacocks that had elaborate trains. Students are likely to experience the same defoliation as Darwin, and so discussing peacock trains provides an ideal opportunity to reinforce how natural pick works, and to introduce the topic of sexual selection.

We began a give-and-take of sexual selection past pointing out that natural pick is oftentimes summarized as "survival of the fittest" only noted that many animals have traits that seem to subtract their chances of survival. Nosotros proposed that the elaborate trains of peacocks are perfect case of such traits; the long feathers would seem to increase the risk of predation. We and then asked our students to come up with as many hypotheses every bit possible to explain why peacocks have such elaborate trains. Students proposed that tail feathers scared away potential predators or helped attract mates. We are not enlightened of whatever inquiry on the power of peacock trains to deter predators, and then we told our students that we were going to restrict our word to the mating preference hypothesis.

Side by side, nosotros showed our class a graph from Petrie's famous mate choice written report at the Whipsnade Zoo (Figure iii in Petrie et al., 1991 ). This graph shows that peacocks with many eyespots (a sign of an elaborate train) garnered more mates than males with fewer eyespots. We as well showed students that peahens mated more often with peacocks with bright tail feathers (Figure 3 in Loyau et al., 2007 ) and mated less oft with males that had eye spots removed by researchers (Effigy iii in Petrie and Halliday, 1994 ). These data respond the question of why peacocks accept elaborate trains: elaborate trains increase their chances of mating. Nosotros explained how such a mating preference could give rise to the elaborate trains using the requirements for natural selection that nosotros used throughout the course (i.eastward., natural selection requires variation, heritability, and differential reproductive success).

The previous word gave rise to a new, interesting question, namely "Why exercise peahens prefer to mate with peacocks with elaborate trains?" We allow our students discuss this for a couple of minutes, solicited answers, discussed possible explanations, and concluded that peacocks with elaborate trains take much amend genes than peacocks with less elaborate trains. Nosotros introduced this to our students equally the "good genes hypothesis" and asked them how they would test information technology. This was a difficult question for our students. Many students suggested examining peacocks with elaborate trains and testing them to determine whether they were healthier or otherwise superior to peacocks with less elaborate trains. The trouble with this experimental design is that both traits (elaborate trains and wellness) are likely influenced past environmental factors. Peacocks raised in favorable weather might take elaborate trains and be salubrious—and not have proficient genes. A rigorous test of the good genes hypothesis must exam whether peacocks with elaborate trains contribute ameliorate genes to offspring than peacocks with less elaborate trains. Petrie (1994) did this by randomly mating peahens and peacocks and showing that the chicks of peacocks with elaborate trains were more likely to survive than the chicks of peacocks with less elaborate trains (run into Figure 2 in Petrie, 1994 ). After we completed this discussion, we formally divers sexual selection, talked nigh why females get to choose, and discussed the consequences of sexual pick.

Evolution of a Circuitous Trait: Antibiotic Resistance in Escherichia coli.

Students frequently have a hard time agreement how complex traits, such as the vertebrate eye, have evolved through a combination of random mutation and natural selection. Darwin (1859) predictable this. He wrote in Chapter half-dozen of the Origin of Species: "To suppose that the eye … could take been formed by natural selection, seems, I freely confess, absurd in the highest possible caste." Darwin and so went on to explain how such circuitous structures could evolve via natural option through the accumulation of small changes, and so long as each pocket-size change improved the power of the organism to survive and reproduce. This is an important concept for students to understand.

We began our discussion of the evolution of circuitous traits by introducing our students to antibiotic resistance in E. coli. E. coli are rod-shaped bacteria that live in the human being digestive tract. Near strains are harmless, but some cause infections that doctors may treat with penicillin or other antibiotics. Some Eastward. coli accept a form of the enzyme β-lactamase that can suspension down the β-lactam band present in penicillin and other antibiotics, including cefotaxime (Baquero and Blazquez, 1997 ). The caste to which E. coli are resistant to antibiotics depends on how chop-chop β-lactamase can interruption down the β-lactam ring of the antibiotic, and this rate depends on the sequence of amino acids in the β-lactamase enzyme. The β-lactamase allele near commonly present in E. coli populations is the TEM-one allele, which provides a small-scale amount of resistance to the antibiotic cefotaxime. In contrast, the TEM-52 allele provides more than 4000 times as much resistance to cefotaxime. TEM-52 differs from TEM-1 by three amino acid substitutions.

In one case nosotros introduced our students to this case study, we asked them "What does the mechanism of natural selection predict almost the evolution of TEM-i β-lactamase to TEM-52 β-lactamase?" Nosotros permit our students hash out this question in pairs, and then randomly called on students for answers. The answer to the question is that TEM-1 must evolve into TEM-52 via three mutations and each mutation must increase the ability of E. coli to survive and reproduce. Our students have not had a hard time deducing this. Weinreich et al. (2006) synthesized East. coli with each possible combination of amino acids at the three sites that needed to be changed and showed that there were indeed sequences of mutations that could change TEM-one to TEM-52, while increasing antibiotic resistance in each stride. We completed this discussion by telling students that this belongings of β-lactamase development—that each mutation must increase fettle—is a general feature of evolution, and the lecture connected with a discussion of the evolution of the vertebrate eye.

Evolution of Behavior: Apparent Suicide in Lemmings.

A common misconception amongst students is that natural selection favors traits that are "practiced for the species." The lemming suicide myth provides an ideal opportunity to discuss why this is not always true and to testify students that the evolutionary origins of donating behaviors require special explanation. We began our give-and-take of lemming suicide past telling students we were going to show them a nature documentary about lemmings, and after the film, they would be asked to answer the question: "Could suicide be an adaptation in lemmings?" Then we showed the grade a three.v-min flick prune from the 1958 Walt Disney documentary White Wilderness. The picture show clip is available on YouTube (www.youtube.com/watch?v=xMZlr5Gf9yY or search world wide web.Youtube.com for "White Wilderness"). The pic appears to show lemmings jumping off cliffs into the Arctic Ocean and swimming to their death. The narrator does non explicitly claim the lemmings are committing suicide, just strongly implies this is what is happening, and most viewers will believe this is what they are watching.

After watching the film, we asked our students "Could suicide be an adaptation in lemmings to preclude overcrowding?" and permit them discuss this in small groups. And then we randomly called on students and discussed the answers nosotros obtained. Through discussion, we attempted to explain that natural selection is unlikely to favor suicide, because individuals that commit suicide will non pass on their genes. Gary Larson's Far Side cartoon showing a bunch of lemmings rushing into the h2o, including 1 wearing a life preserver, was useful for showing how selfish lemmings who did not commit suicide would pass on their genes and cause the frequency of selfish behavior to increment.

Our students seem to quickly understand why self-subversive behavior is unlikely to evolve via natural choice, but struggled to explicate the apparent mass suicide they witnessed in the documentary. For example, some students proposed the lemmings were committing suicide after they had reproduced or because they had a disease they did non want to transmit to their offspring. Others suggested that the lemmings were leaving overcrowded locations in search of a meliorate place to live and reproduce and happened to come beyond a trunk of water in their way. These are reasonable suggestions, simply plow out to be wrong. The bodily explanation reveals more near human being nature than natural selection. According to a documentary aired by the Canadian Broadcasting Company on May 5, 1982 (www.cbc.ca/fifth/cruelcamera/video2.html), the dramatic footage in the pic White Wilderness of lemmings jumping into the Arctic Body of water was faked. Evidently, the lemmings were pushed off of a cliff into a river. And nevertheless, the moving picture won the 1958 Academy Award for all-time documentary.

Assessment

Nosotros used two instruments to mensurate how well students understood natural selection before, during, and after our serial of six exercises. The offset instrument was a 10-question version of the Conceptual Inventory of Natural Option (CINS-abbr; Anderson et al., 2002 ; Fisher, Williams, Lineback, and Anderson, personal communication ; see Table iii below). This is a multiple-pick exam with distracters designed to appeal to students having common misconceptions regarding natural selection and related concepts. In improver, we used seven short essay questions (Table two) to give students an opportunity to answer evolutionary questions in their ain words. The questions nosotros used (Table 2) were variations of questions developed by Bishop and Anderson (1990) for their Open Response Instrument and are similar in form to questions recently described by Nehm et al. (2012) . Each question asked students to explicate how an accommodation in a familiar creature might have evolved. Vi of the seven questions involved the gain of a trait, and one involved the loss of a trait.

Table 2.

Short essay questions used to assess student agreement of natural selection

Cheetahs are able to run faster than threescore miles per hour when chasing prey. How would a biologist explain how the power to run this fast evolved in cheetahs, assuming their ancestors could merely run 20 miles per hour?

Polar bears have white fur that blends in well with their snowy surround. This helps polar bears stalk and hunt seals. Polar bears are believed to take evolved from bears that had brown fur. How would a biologist explain how the white fur of polar bears evolved from bears with brown fur?

Musk oxen are large animals that that live in the coldest parts of the Chill and look something similar shaggy cows. Musk oxen accept the warmest wool of any mammal. How would a biologist explain how musk oxen evolved this warm wool, bold that their ancestors had wool that was less warm?

Flight squirrels accept folds of skin between their front and back legs that allow them to glide (although not wing) betwixt copse. How would a biologist explain how flying squirrels evolved these folds of skin, assuming their ancestors did not take these folds?

Eagles have swell eyesight that allows them to spot mice and other casualty while soaring loftier to a higher place the ground. How would a biologist explain how eagles evolved their neat eye sight, assuming their ancestors had less keen eyesight?

Camels shop fat in their humps, which allows them to travel for long distances without eating. How would a biologist explicate how camels evolved their humps, assuming their ancestors did not accept humps?

Whales are large mammals with streamlined bodies that allow them to swim hands in the ocean. Unlike most mammals, whales do not have hind limbs. How would a biologist explain how whales lost their hind limbs, assuming their ancestors had hind limbs?

Tabular array three.

Frequency of the well-nigh normally selected incorrect answer in the CINS-abbr before (PRE) and afterwards pedagogy (Postal service) teaching in 2011

	Frequency
Question and most popular wrong respond before educational activity	PRE	POST
1. One time a population of finches has lived on a particular island for many years in a relatively steady climate	0.21	0.06
a. The population size continues to grow chop-chop, at maximum rates.
2. What is the all-time manner to characterize the evolutionary changes that occur in a finch population?	0.32	0.08
d. The surroundings causes specific mutations in individual finches to help them survive and reproduce.
3. Imagine that the ancestors of the cactus finch colonized a new island. They did not accept the beak type seen in the cactus finch today. How did the cactus finches' unique bill blazon first ascend?	0.xv	0.08
c. In the descendants of the original birds, the environs gradually caused the genetic changes that were necessary to live on that island.
4. What type of variation in a finch population is passed to the side by side generation?	0.35	0.08
d. Any characteristics that were positively influenced past the environment.
5. What caused populations of finches having dissimilar beak shapes and sizes to get singled-out species?	0.fifty	0.00***
d. The environment of each island gradually molded beak shape in the new species, because that particular shape was needed to obtain the food.
6. A typical natural population consists of hundreds of guppies Which statement best describes the guppies of a single species in an isolated population?	0.41	0.22
b. The guppies share all of the essential characteristics of the species; the minor variations they possess do not bear upon survival or reproduction.
vii. Once a population of guppies has been established for a number of years in a swimming with other organisms, including predators, what will probable happen to the population size, assuming that conditions remain relatively constant.	0.12	0.06
c. The guppy population size will gradually subtract.
8. What is the best way to narrate the evolutionary changes that occur in a guppy population over time?	0.35	0.00***
d. Mutations occur to meet the changing needs of the guppies, considering the environment changes.
nine. Where did variation in spot colors and patterns in guppy populations come up from?	0.24	0.08
a. The guppies needed item spots to survive and reproduce, so those spots developed.
10. What could cause i guppy species to alter into iii species over fourth dimension?	0.47	0.06***
c. Dissimilar mutations in each surroundings occurred, because they were needed, and each population gradually became a new species.

Students were required to complete the CINS-abbr and the short essay questions, but the accuracy of their responses did not affect their grades. Students were told, nonetheless, that their responses would be evaluated equally function of their course participation grade—which comprised two% of their grade for the course.

We used the CINS-abbr to measure how much students learned during our entire unit of measurement on natural selection. We administered the test twice: in the class period before nosotros began our unit of measurement on natural selection and in the class menses after we completed the final of our six learning exercises. We calculated the average score on the CINS-abbr before and later instruction and used this to summate the normalized proceeds (Hake, 1998 ) for the course. We did this in 2011 and 2012. We tested the statistical significance of increases in examination scores using a nonparametric sign test. Scores from students who did not have both the pre- and postinstruction tests were not included in the analysis.

We used vii short essay questions (Table two) to assess learning in two ways. Offset, we used the essay questions as an culling to the CINS-abbr to measure out how much students learned in the entire unit on natural choice. Second, we used these essay questions to monitor learning throughout our vi-lecture unit on natural pick.

Nosotros administered the essay questions before, during, and afterward our unit of measurement on natural selection as follows. Each student answered 1 short essay question earlier nosotros began our unit on natural pick and then one question after each of the six classroom exercises. We were concerned the questions might vary in difficulty, so we randomly divided our class into vii groups and gave each grouping a different question each day we tested the class. Group one answered question one on the first day (before teaching), question ii on the 2d day, so on, answering question seven after the last of the learning exercises. Group two answered question two on the first day, question three on the 2nd day, and question 1 after instruction was over, and and so on. With this design, each student answered a unlike question each day, only the class equally a whole e'er answered the same set of questions. This allowed us to compare the boilerplate score of the class on different days. Administering these short essay questions took a substantial amount of form time, so we did this just in 2011.

Nosotros graded all of the brusque essay questions using the rubric of Andrews et al. (2011b) . This rubric evaluated student responses according to how well they addressed the three primal concepts we emphasized in lecture: variation for a trait within a population, heritability of the trait, and differential reproductive success. We did not hash out the questions or student answers to the questions during grade, and nosotros did non read whatever of the student answers until after the course was over. Nosotros scored student responses in a random order with the name of the student and the date obscured. All of the responses were independently scored by two of the authors (South.T.Chiliad. and T.M.A.), and whatsoever differences in scores were resolved through give-and-take.

We used the short essay data and an analysis of variance (ANOVA) to guess how much students learned during the first lesson. To do this, we compared the average exam score nerveless before education (PRE) with the average score after the first lesson (dog convenance), using ANOVA. We explicitly paired scores for each individual pupil and accounted for potential differences in difficulty among the various questions administered. We also evaluated an interaction term (i.e., = difference between PRE and the first lesson × essay question) to determine whether the degree of initial learning gains differed based on the essay question assigned; we removed the interaction if it was not statistically pregnant.

Next, we were interested in determining whether and the degree to which educatee scores on the essay questions increased throughout our unit of measurement. In this 2d assay, we analyzed only the data nerveless later the showtime lesson (dog convenance) and used a linear regression to quantify the degree of learning gain. Nosotros did not include the exam scores collected before teaching (PRE), considering nosotros wanted to come across whether students benefited from continued instruction (i.e., represented by a nonzero and increasing slope of the line) or whether the gains afterward initial educational activity (which should be highest) were minimal (i.e., represented by a flat line). Nosotros analyzed these data with a generalized linear mixed model that allowed us to account for repeated observations from the aforementioned students by including a compound symmetric covariance structure (Littell et al., 2006 ). We as well accounted for potential differences in difficulty amongst the various questions administered. Additionally, nosotros evaluated an interaction term (i.eastward., = lesson × essay question) to make up one's mind whether changes in the average score over the series of exercises in the unit differed based on the essay question assigned; we removed the interaction if it was non statistically meaning.

Nosotros also used surveys to assess student attitudes toward the six learning activities (run across Table 4 beneath). Participation in these surveys was voluntary and anonymous.

Table 4.

Average responses of surveys used to mensurate student attitudes toward six learning exercises^a

	This action …
	held my interest	challenged me intellectually	was under-standable to me	was a valuable learning feel
Dogs	iv.9	four.5	5.5	v.1
Mice	four.half dozen	four.1	5.4	iv.vii
Humans	4.8	four.6	five.4	v.0
E. coli	4.8	4.6	5.vi	5.0
Peacocks	—	—	—	—
Lemmings	v.4	5.5	5.3	v.2

RESULTS

Information from the CINS-abbr showed that, before instruction, many students appeared to have misconceptions regarding natural selection (Table iii). Between 25 and 50% of our form seemed to believe evolution was caused by the environment changing individuals or species evolving out of demand (Table 3). As we discussed in the Introduction, these are common misconceptions among introductory biology students.

Our class showed impressive learning gains on the x-question version of the CINS both years we administered this test (Figure one). In 2011, pre- and postinstruction CINS-abbr scores were available for 32 of the 41 students in the course. Before instruction, the class boilerplate on the CINS-abbr was 6 (out of 10; σ = two.55). Later on instruction, the boilerplate was 8.88 (σ = 1.45). This corresponds to a normalized gain of 0.72. Analysis of student responses on specific questions suggested the frequency of specific misconceptions declined after instruction (Table 3). In particular, fewer students seemed to have the misconception that development happens considering the surroundings changes individuals.

Pre- and postinstruction test scores on the CINS-abbr in 2011 and 2012.

We observed similar learning gains on the CINS-abbr in 2012. In 2012, pre- and postinstruction CINS-abbr scores were available for 42 of the 47 students in the courses. The average score amid these students increased from 7.14 (σ = 2.55) to 9.24 (σ = i.03) which corresponds to a normalized gain of 0.73. As we discuss beneath, these are exceptionally large learning gains for this test. Notice too that the normalized gain was essentially the aforementioned both years (0.72 vs. 0.73), despite the fact the natural selection unit was taught by ii different instructors and that the composition of the student trunk was essentially different in 2011 and 2012.

Nosotros administered the brusque essay questions seven times to our form of 41 students in 2011 and received 236 student responses. This corresponds to a response charge per unit of 236/(41 × 7) = 82%. Every bit indicated higher up, all questions were graded by two researchers; the correlation between scores was initially 0.82 (all differences resolved via word) Seven student responses were culled earlier nosotros began statistical analysis. Six students missed 3 lectures in a row, and nosotros dropped the responses of these students after these absences. This removed vi responses from our information. We also dropped one student's response to the concluding question. Information technology was the merely response on the last two testing dates that received a score of zippo. The response was unusually short, and the pupil had earned full credit on two previous questions. Furthermore, this response was identified equally an outlier in a general linear model. After dropping these seven responses, we had a final data fix of 229 responses, or a response charge per unit of fourscore%.

Student responses on the curt essay questions showed substantial learning gains. Before instruction, the most common score in the class was 0 points (out of 6) and the average score in the class was ii.84. After instruction, the about common score in the class was 6 points, and the average score was 4.90. This corresponds to a normalized proceeds of 0.65, which, every bit we discuss beneath, compares favorably with results from other courses for similar questions.

On the footing of our assay of student responses to the short essay questions, we quantified a large increment in understanding afterwards the first learning exercise (i.e., difference between PRE and the Domestic dog lesson, Figure 2a). Average scores increased by one.23 points (95% CI = 0.48–1.98) afterward the starting time learning exercise (F(ane, xxx) = eleven.xviii, p = 0.002; difference between PRE and Dog lesson in Effigy 2).

Average scores on the brusk essay questions before instruction (PRE) and afterwards each lesson (Dogs, Mice, etc.) in 2011. (a) The boilerplate score in the course; (b) the average score for each component of the full score included in the rubric. We analyzed the data in (a) in two means. Offset, we quantified the initial learning gains later on one lesson (difference between PRE and Dogs). Next, we determined whether learning gains connected to occur later on initial instruction; the line in (a) shows the predicted increase in the average score for the class throughout the set of exercises. Test scores earlier pedagogy (PRE) were not included in this second analysis.

We also observed that student scores continued to improve throughout the series of our exercises by an average of 0.15 points (95% CI = 0.05–0.25) with each additional practice (F(1, 145) = 9.02, p = 0.003; the Canis familiaris lesson through the Lemmings lesson on Figure 2a), which amounts to an average full increase of 0.75 points (95% CI = 0.25–1.25) over the serial of v exercises. We found picayune bear witness that the management or magnitude of these increases differed based on which essay question students were assigned (i.east., interactive effect = lesson × essay question, initial increase: F(six, 24) = 0.61, p = 0.72; subsequent increment: F(half dozen, 139) = 1.i, p = 0.37).

Our rubric for the curt essay questions contained 3 components: variation, heritability, and differential reproductive success. Student answers consistently received the fewest points for the heritability component (Figure 2b). This component besides seemed to be the most difficult for students to learn. Students' scores for variation and differential reproductive success rapidly rose afterwards the first lesson, and then changed relatively little (especially for the concept of variation). In contrast, scores for the heritability component of student answers steadily increased throughout instruction. This increase was responsible for 77% of the increment in scores subsequently lesson 1.

The survey of student attitudes regarding the half-dozen natural selection exercises showed that students more often than not viewed these lessons positively (Table 4). On average, the class agreed the exercises were interesting, challenging, understandable, and valuable learning experiences. None of the exercises received remarkably low or loftier evaluations, just the lemming discussion (which was the last one taught) did seem to stand out as being viewed as particularly interesting and challenging.

Discussion

We adult and assessed six classroom activities for teaching natural choice to introductory biological science students. All our assessments suggested the activities were both engaging and effective.

The learning gains nosotros observed on the x-question version of the CINS compare very favorably with results from other classrooms. Andrews et al. (2011b) administered the same 10-question CINS-abbr to 33 introductory biology courses randomly sampled from major universities across the United States. Students in these classes took the CINS-abbr before and later instruction on natural selection. The average normalized gain amid these courses was 0.26, and the highest normalized gain observed was 0.68 (T.1000.A., unpublished information). In our classroom, we observed a normalized gain of 0.72 when Due south.T.Grand. taught the form and 0.73 when PhD student taught the lessons. Detect that our learning gains were higher than in any of the other classrooms studied past Andrews et al., even when our unit on natural selection was taught by a graduate student with very footling pedagogy experience. We interpret this as evidence that the lessons we describe here are effective for promoting learning.

Results from assessment using brusque essay questions besides compare favorably with results from other institutions. The normalized gain associated with our six learning activities, equally measured by our short essay questions, was 0.65. The average normalized gain for a short essay question in the national written report of Andrews et al. (2011b) was 0.06, approximately one-10th of what we observed in our classroom. This is additional bear witness that our learning exercises were effective. In that location is, notwithstanding, a methodological difference to note between our assessment and that in the survey of Andrews et al. (2011b) . We used seven brusque essay questions. Andrews et al. (2011b) used but one question: a question on cheetahs very similar to our cheetah question (Table 2). The data, therefore, are not perfectly comparable. However, in that location is no reason to doubtable our set of questions was easier than the chetah question used past Andrews et al. (2011b) . We observed a standardized gain of 0.88 for the chetah question in our classroom; the standardized gains for all of the other questions were lower. Therefore, the set of questions we used to assess our learning exercises was probably harder than the question used past Andrews et al. (2011b) .

Recent research (Nehm and Ha, 2010 ; Nehm et al., 2012 ) has shown the ability of students to answer questions regarding natural pick depends on the context of the question. In detail, Nehm and Ha (2010) showed that questions that involve trait loss, unfamiliar species, or evolution betwixt species are more difficult for students to answer. Had this research been available when we started our project, we would have broadened the telescopic of the short-respond questions in our cess. All the same, we did include one question relating to trait loss (in whales), and this provides us with some insight for how constructive our learning exercises might be for allowing students to reason about natural selection in contexts beyond evolution of a new trait within a species. As expected from Nehm and Ha's (2010) results, scores on our trait-loss question were, on average, lower than the six trait-gain questions. All the same, the learning gains observed with the trait-loss whale questions did not announced to exist lower than those of the other questions. We calculated the normalized gain for each of the vii questions, and the normalized gain for the whale evolution question was 0.60. This was equal to the median of the normalized gain amid our seven questions. Nosotros never discussed or mentioned trait loss at whatever time during the grade, then information technology seems probable that our students learned natural pick well enough that they were able to reason effectively about the relatively novel evolutionary scenario of how traits are lost.

Our pre- and postinstruction testing has shown that our students made impressive progress agreement one of the about difficult concepts in biology. We attribute these learning gains to our classroom exercises and advise four characteristics of our lessons may have been important for promoting learning. Kickoff, we designed these lessons to specifically target pupil misconceptions. We did not merely utilise agile learning—we used specific methods in an active-learning format to help students motion past misconceptions. Second, we explicitly discussed the genetic footing of evolution. Many student misconceptions regarding natural selection relate to genetics, so making a connection to students' prior genetics cognition should be helpful. Tertiary, we presented students with multiple examples of selection at work in a variety of different contexts. Finally, we did our best to create exercises that engaged our students as deeply as possible (Tabular array iv).

Nosotros have attributed the dramatic increase in test scores we observed with our students to the classroom activities nosotros conducted, and accept proposed 4 reasons why our lessons may have been constructive. We do acknowledge, all the same, the design of our study makes information technology impossible to unambiguously aspect student learning to whatsoever specific chemical element of our education. We conducted six classroom activities during our unit on natural pick, but these activities were not the only opportunities students had to acquire. Students were assigned textbook readings before each lesson, and this may have contributed to some of their learning. Similarly, most of our lectures included a off-white amount of ordinary lecturing, and this could accept contributed to student learning. There are many other possibilities. Yet, it is not credible to united states of america that any of these aspects of our course were unique enough to explicate the uniquely loftier learning gains we observed.

Readers may wonder whether whatsoever of our six lessons were more or less effective than others. For example, the average course score on the curt essay questions dropped afterwards the mice and peacock discussions, which raises the question of whether these lessons were effective or not. Unfortunately, our study was not designed to mensurate how much students learned in a specific action. In that location are several reasons why non. First, we did non perform pre- and postinstruction testing for whatsoever of the individual learning activities. We administered the CINS-abbr earlier and after our unit of measurement on natural selection and administered essay questions to the class before, after, and during our unit on natural choice, simply nosotros did non perform any assessment immediately before and/or immediately after whatsoever of the activities described here. Second, our course had only 41 students, and they answered a single essay question each day we did testing during the unit of measurement. This does non requite us a lot of statistical power to measure out how much students learned from 1 lesson to the next. 3rd, student understanding can temporarily "dip" after a lesson, particularly if a lesson presents students with a principle in a new context. When instructors exercise this, students demand to restructure their noesis to deal with the new context (e.g., Bransford and Schwartz, 1999 ; Barnett and Ceci, 2002 ; Lobato, 2008 ). Fourth, it is important to remember that the essay questions provide only one measure of how well students empathise natural selection. Learning gains might have looked different had we used other questions. Finally, we emphasize that our data describes learning gains for the sequence of exercises every bit we taught them. The apparent effectiveness of any lesson almost certainly strongly depends on what is taught before it. For example, the dog-breeding lesson was associated with higher learning gains than the give-and-take of beach mice. If we used the mice lesson commencement and followed it upward with a discussion of dog breeding, we might observe the opposite.

Instructors considering incorporating our learning exercises into their courses may wish to modify them to adjust the specific needs of their courses. If this is done, instructors must ensure these activities give students the opportunity to personally construct an agreement of natural pick. In item, we strongly encourage instructors to brand certain that students: 1) formulate an initial respond to each question on their own, 2) hash out their answers with a few other students, iii) participate in a discussion with the entire class, and 4) recognize how these activities chronicle to the main concepts taught in a course. All of this takes time but is important for learning. There is accumulating evidence that instructors attempting to use active-learning methods practise not include these elements in their instruction, and therefore fail to encounter the learning gains they hope to achieve (Turpen and Finkelstein, 2009 ; Andrews et al., 2011b ; Ebert-May et al., 2011 ).

Instructors using the exercises described in this paper in their classroom should make an effort to assess whether their students are learning natural selection. Some sort of pre- and postinstruction testing is necessary to practise this. The CINS-abbr (Anderson et al., 2002 ), ACORNS (Assessing Contextual Reasoning most Natural Selection) (Nehm et al., 2012 ), or open-response questions (described past Bishop and Anderson, 1990) are reasonable instruments to apply. In that location are no established criteria for what constitutes acceptable learning gains on these tests, simply a normalized gain of ≥0.50 probably represents a substantial achievement.

In the grade we taught, nosotros devoted six 50-min lectures to didactics natural choice. This might seem excessive. Two lines of evidence propose this time was well spent. First, subsequently five lectures on natural choice, many of our students were unable to recognize that suicide was an unlikely adaptation. This implies their understanding of natural choice was still developing or they were unable to depict upon their knowledge when that knowledge was relevant. 2d, the average test score on the curt essay questions we used to appraise understanding of natural selection did non appear to level off. Student answers appeared to be still improving at the end of our series of exercises (Figure 2). In particular, students' understanding of the genetic basis of evolution seemed to be improving (Figure 2).

We will conclude this paper with a few comments on a controversial aspect of moving from traditional lectures to active learning. Most of the learning exercises described hither require a fair amount of time to conduct. This is especially true for the discussions of dog convenance, man evolution, and peacock trains. An instructor using these activities for the first time may find he or she has less time to encompass other material. This could be viewed as a reason not to utilize active learning—specially if instructors experience pressure level to cover as much material as possible. We have two responses. First, the goal of pedagogy is not to cover as much as possible, simply to teach as much as possible, and a large trunk of evidence shows that students acquire more in courses that brand extensive utilize of active learning (e.g., Hake, 1998 ; Knight and Wood, 2005 ; Freeman et al., 2007 ). Second, natural selection is 1 of the nigh important concepts in biology, but it is difficult for students to learn. All evidence suggests many students need specialized and time-consuming instruction to learn natural selection well enough to avoid falling back upon glaringly inaccurate misconceptions. Instructors might encompass more if they used traditional lectures to cover natural pick, merely information technology is hard to imagine what other topics students might learn that would make up for not understanding the principle cause of evolution.

Supplementary Material

ACKNOWLEDGMENTS

Nosotros thank the National Science Foundation for funding (CCLI 0942109) and our students for helping us develop these learning exercises. We thank two bearding reviewers for thoughtful comments that improved the manuscript.

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Which Type Of Selection Would Favor Steadily Increasing Body Size For Wolves In Colder Climates?,

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