Darwin's "imaginary illustrations": creatively teaching evolutionary concepts & the nature of science.
Abstract: An overlooked feature of Darwin's work is his use of "imaginary illustrations" to show that natural selection is competent to produce adaptive, evolutionary change. When set in the context of Darwin's methodology, these thought experiments provide a novel way to teach natural selection and the nature of science.

Key Words: Charles Darwin; natural selection; imaginary illustrations; nature of science.
Article Type: Report
Subject: Teaching (Methods)
Natural selection (Study and teaching)
Biologists (Influence)
Biologists (Works)
Evolution (Study and teaching)
Science (Study and teaching)
Author: Love, Alan C.
Pub Date: 02/01/2010
Publication: Name: The American Biology Teacher Publisher: National Association of Biology Teachers Audience: Academic; Professional Format: Magazine/Journal Subject: Biological sciences; Education Copyright: COPYRIGHT 2010 National Association of Biology Teachers ISSN: 0002-7685
Issue: Date: Feb, 2010 Source Volume: 72 Source Issue: 2
Product: Product Code: 8522101 Biologists; 8520000 Sciences; 8500000 Science, Research & Development NAICS Code: 54171 Research and Development in the Physical, Engineering, and Life Sciences; 5417 Scientific Research and Development Services
Persons: Named Person: Darwin, Charles; Darwin, Charles
Geographic: Geographic Scope: United States Geographic Code: 1USA United States
Accession Number: 245037744
Full Text: [ILLUSTRATION OMITTED]

* Did Darwin Really Say That?

When a new theory or hypothesis is put forward in science, we presume that it will be associated with evidence. This is a hallmark of scientific reasoning--theories or hypotheses should be explicitly supported by empirical evidence. It is one of the reasons for our ever increasing confidence in the theory of evolution. From its original presentation to its current incarnations, the supporting empirical evidence has gone from strength to strength, though not equally on all fronts and certainly with a good dose of controversy. Thus, imagine the surprise when readers of Charles Darwin's On the Origin of Species encounter the following passage under the heading "Illustrations of the Action of Natural Selection": "In order to make clear how, as I believe, natural selection acts, I must beg permission to give one or two imaginary illustrations" (Darwin, 1964 [1859]: p. 90). How can "imaginary illustrations" be evidence for the process of natural selection, which is supposed to explain the evolutionary emergence of exquisite adaptations in different species? Every time I introduce undergraduates to On the Origin of Species, this section induces conceptual vertigo. Many of them are perplexed over how this can be a legitimate form of scientific reasoning: maybe Darwin made a mistake or didn't have a choice? As one of my students recently phrased it: "I'm curious about how Darwin used 'reasoning' and invisible thought experiments to explain things.. .don't you always need proof?" Is Darwin really using these imaginary illustrations as evidence for how natural selection acts or is something else going on?

It turns out that answering the question of what Darwin was doing with "imaginary illustrations" illuminates both the meaning of evolutionary concepts and the nature of science. My aim here is to bring this less familiar component of Darwin's reasoning into the context of teaching biology. His imaginary illustrations offer an accessible and novel way to teach about natural selection and scientific reasoning, especially because they demonstrate an underappreciated aspect of scientific theory testing--evaluating explanatory potential with thought experiments. I begin by exhibiting Darwin's use of imaginary illustrations at various places in On the Origin of Species, which displays how they operate as hypothetical reasoning or thought experiments rather than as attempts to confirm his theory with evidence. Next, I show how Darwin's thought experiments are adaptations of a pattern of reasoning used by Charles Lyell that conforms to the philosophical criteria of causal explanation put forward by John Herschel. Having set Darwin's hypothetical reasoning in the context of this methodology, I detail the intriguing ways in which it yields new materials for teaching biology and aligns with the standards articulated in the National Science Education Standards (National Research Council, 1996). In closing, I emphasize how paying attention to Darwin's thought experiments offers a powerful route to teaching the current explanatory power of evolutionary theory.

* Darwin's Reasoning: A Closer Look

The first imaginary illustration deployed by Darwin in On the Origin of Species is about the hunting abilities of wolves (Figure 1):

Let us take the case of the wolf, which preys on various animals, securing some by craft, some by strength, and some by fleetness; and let us suppose that the fleetest prey a deer for instance, had from any change in the country increased in numbers, or that other prey had decreased in numbers, during that season of the year when the wolf is hardest pressed for food. I can under such circumstances see no reason to doubt that the swiftest and slimmest wolves would have the best chance of surviving, and so be preserved or selected,--provided always that they retained strength to master their prey at this or at some other period of the year, when they might be compelled to prey on other animals. I can see no more reason to doubt this, than that man can improve the fleetness of his grey-hounds by careful and methodical selection. (Darwin, 1964: p. 90)

Darwin encourages us to hypothetically consider wolves that capture prey using three distinct traits: craft, strength, and fleetness. What would happen if prey caught by fleetness increased or prey caught by craft or strength decreased? The swifter and sleeker wolves will be preserved and proliferate, other things being equal ("no reason to doubt"). This would occur just as readily as when breeders select for increased fleetness in their racing dogs, and in the latter case we have an abundance of evidence demonstrating this differential survival and reproduction. Darwin has portrayed the consequences of variation and the struggle for existence in a readily digestible format, but what about heredity? That comes next:

Even without any change in the proportional numbers of the animals on which our wolf preyed, a cub might be born with an innate tendency to pursue certain kinds of prey..[W]e often observe great differences in the natural tendencies of our domestic animals.. The tendency to catch rats rather than mice is known to be inherited. Now, if any slight innate change of habit or of structure benefited an individual wolf, it would have the best chance of surviving and of leaving offspring. Some of its young would probably inherit the same habits or structure, and by the repetition of this process, a new variety might be formed which would either supplant or coexist with the parent-form of wolf. (Darwin, 1964: p. 91)

[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

Darwin's hypothetical scenario represents what natural selection is capable of doing (forming a new variety) when its three necessary conditions are met: there is variation, the variation exhibits an effect on fitness, and the variation is heritable. It is not a description of natural selection in the wild because evidence for the three conditions is from anecdotes and analogies to domesticated contexts. For example, support for innate prey preferences comes from individual testimony: "one cat, according to Mr. St. John, bringing home winged game, another hares or rabbits, and another hunting on marshy ground and almost nightly catching woodcocks or snipes" (p. 91). These anecdotes are littered throughout On the Origin of Species. Some are explicitly analogical in that observed patterns of variation and inheritance in artificial breeding situations are inferred to be applicable to situations in nature.

Darwin proceeds to work through a second case that involves co-adaptation between plant pollination and insect foraging (Figures 2, 3, and 4; Darwin, 1964: pp. 91-95). Although this is still "imaginary," it is obvious that Darwin relies upon observations of insects and plants to argue for the gradual modification of insect foraging behavior and plant pollination mechanisms into a co-adapted suite of characters. Throughout the imaginary case, he repeatedly stresses the three conditions for natural selection. Darwin even avoids getting sidetracked with actual evidence of insect pollination in order to tease out further possibilities in the imaginary illustration, such as diversification through the advantageous separation of male and female reproductive organs in flowers. Bypassing empirical evidence ("I could give many facts"), he produces a hypothetical account of how mouthparts related to feeding could become tailored for procuring nectar from particular flower morphologies. The three conditions for natural selection are specifically noted (cf. Figure 4): variation ("accidental deviation ... in the curvature and length of the proboscis"), fitness effects ("would be able to obtain its food more quickly, and so have a better chance of living and leaving descendants"), and heritability ("descendants would probably inherit a tendency to a similar slight deviation of structure").

[FIGURE 3 OMITTED]

Darwin does not conclude from these imaginary illustrations that they provide empirical confirmation of his theory. Rather, they make it more plausible and intelligible.

Thus I can understand how a flower and a bee might slowly become, either simultaneously or one after the other, modified and adapted in the most perfect manner to each other, by the continued preservation of individuals presenting mutual and slightly favourable deviations of structure. (Darwin, 1964: p. 95)

That he emphasizes plausibility rather than confirmation should make it unsurprising that Darwin also uses imaginary illustrations when trying to blunt objections to natural selection producing significant transitions between structures that are adapted to specific environments. Without clear evidence, fossil or otherwise, it is critical to have plausibility for cases such as the origin of whales, where structures originally adapted to terrestrial environments now serve aquatic habits.

Darwin opts for a fictional case to establish the conceivability of natural selection bringing about these transitions. Given the existence of terrestrial species within predominantly "water-breathing classes" and a trait appearing in diverse groups ("flying in birds and mammals"), "it is conceivable that flying-fish, which now glide far through the air, slightly rising and turning by the aid of their fluttering fins, might have been modified into perfectly winged animals" (Darwin, 1964: p. 182). Darwin's hypothetical reasoning about transitions also yields an argument about what to expect in the fossil record:

The linkage of reasoning about the imperfection of the fossil record and the plausibility of selection producing major transitions of structure reappears with another imaginary illustration in chapter 9 ("On the Imperfection of the Geological Record"), which also addresses the objection about the rarity of "numerous, fine, intermediate varieties" (p. 299). Darwin reasons hypothetically about what we should plausibly expect in the fossil record:

* Adapting Lyell's Reasoning: Imaginary Illustrations as Thought Experiments

Darwin didn't make a mistake when he used hypothetical reasoning. It is clearly a recurring feature of On the Origin of Species. We might be tempted to think that he was forced by the circumstance of hasty publication to adopt them. This would explain why Darwin's critics seized on this aspect of his reasoning. Richard Owen (1860: p. 516) singled out Darwin's use of imaginary illustrations and highlighted their evidential deficit:

If only Darwin had had more time available before publishing, he would have replaced the imaginary illustrations with empirical evidence that supported his conclusions--unfortunately, it's just not true.

If we examine the massive Natural Selection (the "big species book"), which Darwin abandoned to write the "abstract" we now know as On the Origin of Species (in response to receiving Alfred Russel Wallace's paper on species origination), we find that Darwin had already adopted imaginary illustrations in discussing both the operation of natural selection and the intelligibility of evolutionary transitions (Stauffer, 1975). Darwin had employed hypothetical reasoning for at least two decades, in both his geological writings and his notebook theorizing (Lennox, 2005). He also does not remove it from subsequent editions of On the Origin of Species, even though he made other changes in response to criticisms, including specific modifications to imaginary illustrations in response to Fleming Jenkin's objections regarding the frequency of beneficial variations (Lennox, 1991). Both the wolf and insect-pollination examples reappear in all six editions (Peckham, 1959).

The source of Darwin's hypothetical reasoning goes a long way toward explaining its central role in his methodology. Darwin was following in the footsteps of Lyell's geological reasoning (Herbert, 2005). A clue to Darwin's method is found shortly after his first two imaginary illustrations:

I am well aware that this doctrine of natural section, exemplified in the above imaginary instances, is open to the same objections which were at first urged against Sir Charles Lyell's noble views on the "the modern changes of the earth, as illustrative of geology;" but we now very seldom hear the action, for instance, of the coastwaves, called a trifling and insignificant cause, when applied to the excavation of gigantic valleys or to the formation of the longest lines of inland cliffs. (Darwin, 1964: pp. 95-96)

The phrase "doctrine of natural selection" has an odd ring to our ears but is intended to encapsulate both that natural selection operates and that it is competent to produce the adaptive changes that lead to new species. Darwin sets up a parallel between the objections to Lyellian geological causes being "trifling and insignificant" in the production of large-scale effects through uniform but regular operation, and objections to natural selection accumulating small, heritable, beneficial variations to produce new species and complex adaptations. Darwin's hypothetical reasoning is an adaptation (or co-opting) of a strategy routinely used by Lyell. For example, in the second volume of his Principles of Geology, Lyell (1991 [1832]) offered numerous imaginary illustrations (though he didn't call them this) to show how various means of dispersal could account for the geographic ranges of different species (Lennox, 2005). The key difference is that Darwin was self-aware of this mode of reasoning.

Darwin's imaginary illustrations, like laboratory experiments, are utilized for testing his theory, but they are not meant to test its truth ("empirical confirmation"). Rather, they test its explanatory potential ("understanding" or "intelligibility") as thought experiments (Lennox, 1991). Two tests of explanatory potential are visible in Darwin's thought experiments: (1) establishing that natural selection is competent to produce adaptive, evolutionary change generally; and (2) establishing that natural selection is competent to produce adaptive, evolutionary change in specific problem cases. The former appear in chapter 4 when Darwin treats the action of natural selection, and the latter appear in discussions of difficulties, such as evolutionary transitions or fossil-record imperfection (chapters 6-9). Thus, counter to our initial intuitions, imaginary illustrations are neither mistakes nor foisted upon Darwin by circumstance. They are a genuine part of his method of theory evaluation, but not in the way usually emphasized when discussing the testing of scientific theories.

A rationale for Darwin's method is discernible when we focus on the background to Lyell's geological reasoning. Lyell was attempting to exemplify a scientific standard for causal explanations of historical events advocated by his colleague John Herschel. An inductive methodology requires the identification of real causes operating in nature and a demonstration that they are "competent, under different modifications, to the production of a great multitude of effects" (Herschel, 1987 [1830]: p. 144). This can be codified into three criteria of demonstration: (1) that a cause exists in nature, (2) that a cause is competent to produce effects, and (3) that a cause is responsible for producing particular effects. There are a number of reasons to think that this tripartite standard animates On the Origin of Species (Hodge, 1992). Therefore, the parallel in reasoning about past geological and biological phenomena is attributable to a common cause--a commitment to what counts as a good scientific explanation. In particular, Darwin is using thought experiments to persuade us of the second criterion, to establish that natural selection is competent to produce adaptive, evolutionary change both generally and in specific problem cases.

This interpretation of Darwin's methodology is confirmed by the fact that he also employs thought experiments when he introduces sexual selection. The strategy was part and parcel of Darwin's argument for how selection processes are capable of producing adaptive effects over long periods. When he introduces the principle of why advantageous variations that occur at or near maturity are preserved, we hear a familiar refrain: "As this principle has an important bearing on sexual selection, it may be advisable to give an imaginary illustration" (Darwin, 1981 [1871], vol. 1: p. 298). The conclusion is not an empirical confirmation of sexual selection but a defense of its competence to produce the requisite effects:

And, just as in On the Origin of Species, Darwin deployed his hypothetical reasoning when the situation was the most perplexing: "An imaginary illustration will best aid us in seeing the difficulty of the case" (Darwin, 1981, vol. 2: p. 155). He readily averred to useful fictions to demonstrate the competence of natural selection as a cause of adaptive evolutionary change, both generally and for problematic cases.

* Putting It to Use Pedagogically

Although Darwin's methodology is intrinsically interesting, our present rationale for scrutinizing his imaginary illustrations is their pedagogical value. The first feature to highlight is the availability and affordability of these materials. Little adjustment is required to use them, regardless of decisions about textbook adoption. No significant cost is incurred, because the materials are freely available online (http://darwin-online. org.uk/). In addition, these readings are accessible to a wide range of students. They can be used at different levels of instruction, both because the excerpts are relatively short and because they do not assume significant background knowledge about the topic. In light of this, they constitute a fitting introduction to discussions of evolution by natural selection.

Beyond availability, affordability, and accessibility, Darwin's treatment of natural selection in terms of thought experiments is pedagogically appropriate because it aligns with numerous aspects of the National Science Education Standards (National Research Council, 1996). Students who understand how Darwin's thought experiments work are able to "engage intelligently in public discourse and debate matters of scientific and technological concern" (National Research Council, 1996: p. 13) for two main reasons. First, Darwin's hypothetical reasoning provides an accessible introduction to the concept of natural selection that draws attention to the three conditions necessary for its operation: variation, fitness, and inheritance. Second, Darwin's thought experiments encourage students to grapple with the diverse nature of scientific reasoning and how theories mature over time (Farber, 2003), which is also important for educators teaching evolution (Rutledge & Warden, 2000). A better understanding of how natural selection operates and how scientific reasoning works undergirds the scientifically literate populace envisioned in the national standards and can be observed when set alongside specific teacher and content standards.

One of the first teacher standards for science education concerns choosing what and how to teach: "Select science content and adapt and design curricula to meet the interests, knowledge, understanding, abilities, and experiences of students" (National Research Council, 1996: p. 30). Darwin's thought experiments allow each of these to be met explicitly and encourage educators to "orchestrate discourse among students about scientific ideas" (p. 32). They rely on very little background knowledge of biology and, therefore, are appropriate for most students. They can directly connect with student interest and experience by being reworked using local flora or fauna. Darwin's thought experiments also do not require sophisticated reasoning abilities (i.e., their logical structure is not complex). As a consequence, they facilitate "discussion based on a shared understanding of rules of scientific discourse" (p. 46) because the spotlight is not on the biological facts but on the structure of Darwin's argument.

The value of thought experiments only increases when we turn to the content standards. In the life-science standards for grades 5-8, Darwin's thought experiments stimulate reflection on the causes of diversity and adaptations and draw attention to the significance of reproduction and heredity for natural selection. For grades 9-12, they are an ideal introduction to the concept of natural selection and its explanatory role in the theory of evolution. This includes the production of complex adaptations and fossil record patterns, which are acknowledged stumbling blocks for students:

Darwin's thought experiments are aimed directly at making this conceptual connection. They also enable a dissection of the details of the process of evolution by natural selection because they accentuate its three necessary conditions: variation, fitness, and inheritance (see Figure 4). Alters and Nelson (2002) argued that the inability of students to distinguish between the separate processes involved in evolution by natural selection is a fundamental and widespread misconception. Other misconceptions are also naturally addressed via Darwin's thought experiments, such as (1) viewing natural selection as operating on whole species rather than on the differential effects of heritable variation in individuals or (2) confusion about population changes that result from differential reproduction rather than from individuals in a population changing. One can move fluidly from the qualitative expression of natural selection in Darwin's thought experiments to a quantitative formulation in terms of modern population genetics or more extended project-based units that cover further aspects of the content standards (Cook, 2009). Darwin's thought experiments also highlight the interdependence of organisms, in that he chose interspecific selection processes, whether antagonistic (wolves preying on deer) or synergistic (insect foraging and plant pollination).

Finally, Darwin's hypothetical reasoning introduces material relevant to the history and nature of science content standards. They are a compact illustration of methodological naturalism:

[A]s modern geology has almost banished such views as the excavation of a great valley by a single diluvial wave, so will natural selection, if it be a true principle, banish the belief of the continued creation of new organic beings. (Darwin, 1964: p. 96)

Darwin's thought experiments offer accessible historical vignettes from 19th-century scientific investigation, shedding light on the dynamics of controversy and contrasting it with our current situation (Figure 4). They also reveal that theory evaluation involves more than empirical confirmation and remind students of the interplay between the formulation and evaluation of explanations. In this regard, Darwin's thought experiments address fundamental concepts that underlie the content standards, such as "Evidence, Models, and Explanation" (National Research Council, 1996: chapter 6) and make aspects of inquiry concrete (e.g., proposing possible explanations). All of these features together exemplify the changing emphases in the standards. Thought experiments highlight understanding concepts rather than knowing facts; they emphasize fundamental concepts rather than covering many topics; and they stress science as argument and explanation instead of science as exploration and experiment.

* Cause for Celebration?

By this point a worry may have occurred to the reader. When Darwin compared his reasoning about natural selection to Lyell's reasoning about geological causes, he opened himself up to an objection. Lyell and other geologists (including Darwin; see Herbert, 2005) had collected a wide array of data in order to argue for the competence and responsibility of geological causes. Darwin did not have corresponding measurements of natural selection in the wild, which implies that "responsibility" was out of reach in On the Origin of Species. Doesn't this reveal an embarrassing weakness in the theory of evolution by natural selection? Not at all. In fact, this worry about Darwin's thought experiments can be transformed into an increased confidence in contemporary evolutionary theory.

Was it impossible for Darwin to collect data on natural selection in the wild? Maybe, but the function of his imaginary illustrations remains intact. They were not intended as empirical confirmation but as an exhibition of explanatory potential. We now have updated thought experiments that show with a high degree of sophistication just how plausible it is for a complex structure to originate by natural selection (Nilsson & Pelger, 1994). Even more importantly, Darwin's thought experiments show how evolutionary theory has matured over the past 150 years. This is another aspect of the nature of science--theories develop and progress over time. We have a host of documented examples of natural selection operating in the wild with all the data an avid Herschelian could desire (Endler, 1986). We also have meticulously detailed examples of the evolution of co-adapted traits like hormones and receptors (Dean & Thornton, 2007). Where Darwin introduced thought experiments to demonstrate the competency of natural selection, we now have in hand demonstrations of its responsibility for the origin of species and evolution of traits (http://www.nature.com/evolutiongems). Darwin's methodology provides an ideal basis for teaching evolution by natural selection because with contemporary biology we can directly answer Owen's (1860) criticism. Natural selection is not "just one of those obvious possibilities that might float through the imagination of any speculative naturalist" but instead is genuinely "explanatory of the origin of species." It is difficult to think of a better anniversary gift: Darwin's illustrations are no longer imaginary.

* Acknowledgments

I am grateful to two anonymous referees for providing constructive feedback and helpful suggestions on the manuscript. Thanks also to Olle Pellmyr for the permission to use Figure 2 and to Mark Borrello for consultation on Figure 4.

DOI: 10.1525/abt.2010.72.2.6

References

Alters, B.J. & Nelson, C.E. (2002). Perspective: teaching evolution in higher education. Evolution, 56, 1891-1901.

Cook, K. (2009). A suggested project-based evolution unit for high school: teaching content through application. American Biology Teacher, 71, 95-98.

Darwin, C. (1964). On the Origin of Species. [Reprint of 1859 edition.] Cambridge, MA: Harvard University Press.

Darwin, C. (1981). The Descent of Man and Selection in Relation to Sex. [Reprint of 1871 edition.] Princeton, NJ: Princeton University Press.

Dean, A.M. & Thornton, J.W. (2007). Mechanistic approaches to the study of evolution: the functional synthesis. Nature Reviews Genetics, 8, 675-688.

Endler, J.A. (1986). Natural Selection in the Wild. Princeton, NJ: Princeton University Press.

Farber, P. (2003). Teaching evolution & the nature of science. American Biology Teacher, 65, 347-354.

Herbert, S. (2005). Charles Darwin, Geologist. Ithaca, NY: Cornell University Press.

Herschel, J.F.W. (1987). A Preliminary Discourse on the Study of Natural Philosophy. [Reprint of 1830 edition.] Chicago, IL: University of Chicago Press.

Hodge, M.J.S. (1992). Discussion: Darwin's argument in the Origin. Philosophy of Science, 59, 461-464.

Lennox, J.G. (1991). Darwinian thought experiments: a function for just so stories. In T. Horowitz & G.J. Massey (Eds.), Thought Experiments in Science and Philosophy (pp. 223-245). Savage, MD: Rowman and Littlefield.

Lennox, J.G. (2005). Darwin's methodological evolution. Journal of the History of Biology, 38, 85-99.

Lyell, C. (1990). Principles of Geology, Vol. 2. [Reprint of 1832 edition.] Chicago, IL: University of Chicago Press.

National Research Council. (1996). National Science Education Standards. Washington, DC: National Academy Press.

Nilsson, D-E. & Pelger, S. (1994). A pessimistic estimate of the time required for an eye to evolve. Proceedings of the Royal Society of London, Series B, 256, 53-58.

[Owen, R.] (1860). [Review of Origin & other works]. Edinburgh Review, 111, 487-532.

Peckham, M., Ed. (1959). The Origin of Species by Charles Darwin: A Variorum Text.

Philadelphia: University of Pennsylvania Press. Rutledge, M.L. & Warden, M.A. (2000). Evolutionary theory, the nature of science

& high school biology teachers: critical relationships. American Biology Teacher, 62, 23-31.

Stauffer, R.C., Ed. (1975). Charles Darwin's Natural Selection, being the second part of his big species book written from 1856 to 1858. Cambridge, MA: Cambridge University Press.

ALAN C. LOVE is Assistant Professor of Philosophy at the University of Minnesota, 831 Heller Hall, 271 19th Ave. S, Minneapolis, MN 55455; e-mail: aclove@umn.edu.
Figure 4. Using Darwin's "imaginary illustrations" in the classroom.

The following template uses Darwin's thought experiment of
co-adaptation between plants and insects to stimulate discussion
about the concept of natural selection and the nature of science.
Although it can be incorporated into a variety of lesson plans, it
is very natural to use it early on as a qualitative introduction
that leads to more quantitative approaches. Students first read the
"imaginary illustration" of how plants and insects coevolve (On the
Origin of Species pp. 91-95; http://darwin-online.org.uk/) and then
work through a related set of questions.

The first set of questions (1--1) can be done individually or in
groups. Questions 1-2 focus on the three necessary components of
natural selection and their effects over many generations.
Questions 3--1 focus on the interplay of assumptions in the thought
experiment and evidence that supports them. The next set of
questions (5-6) can be done in a general discussion that utilizes
student answers to the earlier questions. Question 5 focuses on
Darwin's mode of reasoning (exploring explanatory potential rather
than confirming his theory). Question 6 focuses on how thought
experiments facilitate the gathering of data that support the
theory of evolution by natural selection.

The questions are intentionally modular so that educators can
modify and adapt them to their own teaching context (e.g., focus
only on the plant or skip the discussion about assumptions).
Students' answers need not include everything listed below, but the
relevant quotations from Darwin are given as a preparation aid. If
desired, a contemporary reading on plant-insect coevolution can be
introduced to draw a contrast between scientific understanding
during Darwin's time and our own (see below).

Questions

(1) Identify the three necessary components required for natural
selection to operate on the plant and the insect, as given by
Darwin in the imaginary illustration.

Plant

Variation: "Those individual flowers which had the largest glands
or nectaries, and which excreted most nectar, would be oftenest
visited by insects"; "those individuals which produced more and
more pollen, and had larger and larger anthers"

Fitness Effects: "The flowers of two distinct individuals of the
same species would thus get crossed; and the act of crossing, we
have good reason to believe (as will hereafter be more fully
alluded to), would produce very vigorous seedlings, which
consequently would have the best chance of flourishing and
surviving"; "Those flowers, also, which had their stamens and
pistils placed, in relation to the size and habits of the
particular insects which visited them, so as to favour in any
degree the transportal of their pollen from flower to flower"; "yet
if a little pollen were carried, at first occasionally and then
habitually, by the pollen-devouring insects from flower to flower,
and a cross thus effected, although nine-tenths of the pollen were
destroyed, it might still be a great gain to the plant"

Inheritance: "Some of these seedlings would probably inherit the
nectar-excreting power"

Insect

Variation: "an accidental deviation in the size and form of the
body, or in the curvature and length of the proboscis"

Fitness Effects: "certain insects depended in main part on its
nectar for food"; "might profit a bee or other insect, so that an
individual so characterised would be able to obtain its food more
quickly, and so have a better chance of living and leaving
descendants"; "it might be a great advantage to the hive-bee to
have a slightly longer or differently constructed proboscis"

Inheritance: "Its descendants would probably inherit a tendency to
a similar slight deviation of structure"

(2) What does Darwin claim is the result of these three components
occurring over many generations in the case of the plant?

"oftenest visited by insects, and would be oftenest crossed; and so
in the long-run would gain the upper hand"; "Those flowers, also,
which had their stamens and pistils placed, in relation to the size
and habits of the particular insects which visited them, ... would
likewise be favoured or selected"; "those individuals which
produced more and more pollen, and had larger and larger anthers,
would be selected"

(3) What other assumptions are involved in the imaginary
illustration (as given by Darwin)?

(a) Existence of a secreting mechanism: "Certain plants excrete a
sweet juice, apparently for the sake of eliminating something
injurious from their sap"

(b) Location of the secreting mechanism: "Let us now suppose a
little sweet juice or nectar to be excreted by the inner bases of
the petals of a flower"

(c) Flower pollination: "insects in seeking the nectar would get
dusted with pollen, and would certainly often transport the pollen
from one flower to the stigma of another flower"

(d) Content of nectar is desirable: "sweet juice"

(e) Distribution of plants: "we may suppose the plant ... to be a
common plant"

Follow-up question: Do you think these assumptions are reasonable?
Why or why not?

(4) Is there evidence that supports these additional assumptions
(as given by Darwin)?

(a) Yes: "this is effected by glands at the base of the stipules in
some Leguminosa, and at the back of the leaf of the common laurel"

(b) No, not in this passage

(c) Yes: "they would, unintentionally on their part, regularly
carry pollen from flower to flower; and that they can most
effectually do this, I could easily show by many striking
instances. I will give only one" (holly tree example and
experiments)

(d) Yes: "This juice, though small in quantity, is greedily sought
by insects"

(e) No, not in this passage

Follow-up question: Do you think the evidence offered is
persuasive? Why or why not?

(5) Do you think that the thought experiment shows that natural
selection is a plausible account of how adaptations originate? Why
or why not?

Darwin's conclusion: "Thus I can understand how a flower and a bee
might slowly become, either simultaneously or one after the other,
modified and adapted in the most perfect manner to each other, by
the continued preservation of individuals presenting mutual and
slightly favourable deviations of structure."

This part of the discussion should be very open-ended, and answers
usually vary. It is essential to distinguish the two different
styles of reasoning: exploring explanatory potential and confirming
a theory with evidence. The nature of science should become salient
because students who find the thought experiment unconvincing will
often be assuming a particular (faulty) view about how science
works and those who find it convincing will often not be able to
say why. Misunderstandings about the role of variation, fitness
effects, and inheritance in evolution by natural selection usually
surface as well.

(6) How does focusing on the three distinct components in the
thought experiment suggest where to look for evidence for
evolutionary change by natural selection?

Answers will vary, but the critical point is for students to
discern that you need to (minimally) document (a) variation, (b)
fitness effects, and (c) inheritance for the relevant traits in
both plants and insects. Some students may also perceive the need
to do these measurements over multiple generations. A key
conclusion is that the thought experiment is able to guide research
and therefore plays an ongoing role in scientific investigation.
Also, students can discuss how difficult it would be to gather
different kinds of evidence for these components, as well as the
assumptions that Darwin does not give evidence for in this passage.
Students often seize on the fact that Darwin devotes very little of
the thought experiment to illustrating inheritance, as compared
with variation and fitness effects. This can lead to a discussion
of the differences between what Darwin knew and what we know about
variation or as a transition into a lesson on genetic variation.

Optional Supplement

Students read the following recent article about coevolution
between insects and plants before answering the following questions
(either in small groups or in a larger discussion format), which
draws attention to the differences between what Darwin knew and the
increased explanatory power of contemporary evolutionary theory.
This illustrates how theories develop over time, which is another
important aspect of the nature of science.

Sessions, L.A. & Johnson, S.D. (2005). The flower and the fly.
Natural History, 114, 58-63. Questions

(1) In what ways has subsequent research provided evidence for the
explanatory potential that Darwin argued could be found in the
theory of evolution by natural selection?

(2) In what ways has subsequent research gone beyond what Darwin
would have understood (e.g., finding new kinds of evidence, testing
specific hypotheses, or introducing new concepts)?

(3) What kinds of evidence are contemporary biologists looking for
to confirm or extend our understanding of coevolutionary processes?

*Another option is to segue into a discussion of Darwin's work on
orchids, where he fleshed out many of his own ideas about
coevolution in more detail (available at
http://darwin-online.org.uk/). This would be a suitable strategy if
the history of science were being stressed in the unit on natural
selection.

Resources and Further Reading

Darwin, C. (1862). On the Various Contrivances by which British and
Foreign Orchids are Fertilised by Insects, and on the Good Effects
of Intercrossing. London: John Murray.

Oregon Public Radio and PBS. (2001). Co-evolution of Plants and
Pollinators. American Field Guide Teacher Resource Packet.
[Online.] Available at
http://www.pbs.org/americanfieldguide/teachers/insects/insects_sum.html.

Pellmyr, O. (2003). Yuccas, yucca moths, and coevolution: a review.
Annals of the Missouri Botanical Garden, 90, 35-55.

Thompson, J.N. (1994) The Coevolutionary Process. Chicago, IL:
University of Chicago Press.


[T]o return to our imaginary illustration of
   the flying-fish, it does not seem probable
   that fishes capable of true flight would have
   been developed under many subordinate
   forms, ... until their organs of flight had
   come to a high stage of perfection, so as to
   have given them a decided advantage over
   other animals in the battle for life. Hence
   the chance of discovering species with transitional
   grades of structure in a fossil condition
   will always be less, from their having
   existed in lesser numbers, than in the case
   of species with fully developed structures.
   (p. 183)


[W]e have no right to expect to find in our
   geological formations, an infinite number of
   those fine transitional forms.... We ought
   only to look for a few links, some more
   closely, some more distantly related to each
   other. (p. 301)


Now this ... is no very profound or recondite
   surmise; it is just one of those obvious
   possibilities that might float through the
   imagination of any speculative naturalist;
   only, the sober searcher after truth would
   prefer a blameless silence to sending the
   proposition forth as explanatory of the origin
   of species, without its inductive formation.


Thus we can understand how it is that variations
   which must originally have appeared
   rather late in life have alone or in chief part
   been preserved for the development of secondary
   sexual characters; and the remarkable
   coincidence between the periods of
   variability and of sexual selection is intelligible.
   (p. 298)


[S]tudents often do not understand natural
   selection because they fail to make a conceptual
   connection between the occurrence
   of new variations in a population and the
   potential effect of those variations on the
   long-term survival of the species. (National
   Research Council, 1996: p. 184)
Gale Copyright: Copyright 2010 Gale, Cengage Learning. All rights reserved.