CBE Life Sci Educ-2016-Mulnix PDF

ESSAY
STEM Faculty as Learners in Pedagogical

Reform and the Role of Research
Articles as Professional Development
Opportunities
Amy B. Mulnix*
Franklin and Marshall College, Lancaster, PA 17604
ABSTRACT
Discipline-based education research (DBER) publications are opportunities for professional development around science, technology, engineering, and mathematics (STEM) education reform. Learning theory tells us these publications could be more impactful if authors,
reviewers, and editors pay greater attention to linking principles and practice. This approach, which considers faculty as learners and STEM education reform as content, has the
potential to better support faculty members because it promotes a deeper understanding
of the reasons why a pedagogical change is effective. This depth of understanding is necessary for faculty members to successfully transfer new knowledge to their own contexts.
A challenge ahead for the emergent learning sciences is to better integrate findings from
across sister disciplines; DBER reports can take a step in that direction while improving
their usefulness for instructors.
CONTRIBUTIONS OF DISCIPLINE-BASED EDUCATION RESEARCH

PUBLICATIONS TO STEM EDUCATION REFORM
The emerging field of the learning sciences is integrating and extending perspectives
from cognitive, social, and cultural psychology; cognitive neuroscience; education;
discipline-based education research (DBER); and many other fields (e.g., Ambrose
etal., 2010; Slavich and Zimbardo, 2012). These insights promise to deepen learning
for all students and close achievement gaps for groups underrepresented in the sciences (Mulnix etal., 2016). DBER has made and continues to make essential contributions to improving science, technology, engineering, and mathematics (STEM) undergraduate education (National Research Council [NRC], 2012a; Kober, 2015).
Especially in recent years, the appearance of DBER in premier research journals such
as Science (e.g., Haak etal., 2011), Proceedings of the National Academy of Sciences
(e.g., Freeman etal., 2014), and Cell (e.g., Stockwell etal., 2015) has made these
exciting advances in learning sciences apparent to an even broader STEM audience.
But there is potential for DBER publications to contribute more to STEM education
reform than just reporting research findings (Brownell and Tanner, 2012). For
instance, as points of contact with faculty members who are not regular readers of
education literature, articles in high-impact journals can raise awareness and understanding of evidence-based pedagogies (EBP; Bouwma-Gearhart, 2012; Brownell and
Tanner, 2012; Lund and Stains, 2015). Additionally, the relatively high citation rate
and visibility of such articles likely makes them models for subsequent publications.
And while some recent research suggests that lack of knowledge is a diminishing
barrier to reform efforts (Borrego etal., 2010; Henderson etal., 2012; Lund and
Stains, 2015), models (e.g., Table 1) for how change occurs in STEM pedagogy suggest publications have impact at points beyond introduction to ideas (Henderson etal.,
2012; Lund and Stains, 2015). For example, Bouwma-Gearhart (2012) suggests that
the education literature provides resources for faculty members both to explore EBP in
Deborah Allen, Monitoring Editor

Submitted December 8, 2015; Revised April 13,
2016; Accepted April 15, 2016
CBE Life Sci Educ December 1, 2016 15:es8
DOI:10.1187/cbe.15-12-0251
*Address correspondence to: Amy B. Mulnix (amy
.mulnix@fandm.edu).
2016 A. B. Mulnix. CBELife Sciences
Education 2016 The American Society for Cell
Biology. This article is distributed by The
American Society for Cell Biology under license
from the author(s). It is available to the public
under an AttributionNoncommercialShare
Alike 3.0 Unported Creative Commons License
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by-nc-sa/3.0).
ASCB and The American Society for Cell
Biology are registered trademarks of The
American Society for Cell Biology.
Supplemental Material can be found at:

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A. B. Mulnix
TABLE 1. Faculty use of literature during reform of teaching
practices
Innovationdecision model of change
Knowledge vs. no
knowledge
Y/Y
Tried vs. not

tried
Y/Y
Current user
vs. former
user
Y/N
High user vs.

low user
Y/Y
Henderson etal. (2012) identified variables correlated with use of EBP. Reading
the literature regularly was the only significant variable identified as important at
more than two stages in the innovationdecision model. Y/Y indicates that the
regular reading of the literature was significantly correlated as an individual variable and when the other significant variables were controlled for. Y/N indicates
that regular reading of the literature was positively correlated when examined
individually but not when the other variables were controlled for.
low-risk, private environments and to troubleshoot new

approaches in real time.
As shown in Table 1 (Henderson etal. 2012), some research
bears this out: regular reading of STEM education journals does
positively correlate with persistence of physics faculty at multiple stages in the transition to EBP. These stages include knowledge acquisition, motivation to try pedagogical innovations,
and being a high-frequency user. This makes intuitive sense.
Faculty members turn to education literature to initially learn
about EBP and become enthused by the reported positive
results. After gaining experience, they further explore the literature to add more practices to their alternative pedagogy tool
kits (e.g., Major and Palmer, 2006).
Faculty members also encounter educational research literature as part of professional development programs. For example, journal clubs1 (DaRosa and Simpson, 2012; Pusateri, 2014)
and learning communities (McDaniel 1987; Cox, 2001, 2004;
Lenning etal., 2013; Stewart, 2014) are relatively common
opportunities offered by teaching and learning centers. Although
practices vary, both of these activities use publications to
improve pedagogical knowledge and practice. Faculty members
gather to have deep and critical discussions about the findings,
limitations, and implications of research. In some graduate and
postdoctoral programs (e.g., www.cirtl.net/CIRTLReads-2015
-2016; http://otl.wayne.edu/programs/stem.php) learning science research literature is used to strengthen pedagogical
knowledge and, hopefully, teaching skills. Journal clubs, familiar to many STEM faculty for their use in discussion of disciplinary research papers, are typically less demanding and structured than formal learning communities (Cox, 2001, 2004;
Vescio etal., 2008), which include intentional application of
scholarly work in the classroom and an expectation of shared
practice, such as observing one anothers teaching. Additionally, professional developers often send research articles to
interested faculty members, suggest readings for learning communities, provide bibliographies to workshop participants, or
present a seminar based on particularly noteworthy research
findings.
But change is a context-dependent process (Kezar, 2009);
individuals, departments, and institutions differ in how they
approach and accomplish STEM education reform. Lund and
An Internet search identified journal clubs that examine education literature to
be offerings within teaching and learning centers at Yale University, the University of Minnesota, and the Georgia Institute of Technology, among many others.
15:es8, 2
Stains (2015, p. 23) report that DBER articles are largely

underutilized for learning about and implementing EBP by
physics, chemistry, and biology faculty within a single institution. Henderson etal. (2012) also found the correlation of literature use and pedagogical change by STEM faculty to be
complex; although several stages of change are correlated with
use of EBP, continued use after trying new approaches is not
when other variables (e.g., gender, class size) are controlled
for. From this and other data, Henderson etal. (2012, p. 11)
suggest that research articles and other dissemination methods
such as workshops do not support maintained use of these
innovations.
This complexity should not be surprising. Not only is change
context dependent, it is messy even under the best conditions.
It requires iterative practice and is rarely, if ever, accomplished
through reading alone. And, while the primary goal of a
research article is to report findings rather than support STEM
faculty change, pedagogical reform is progressing slowly
(American Association of Colleges and Universities [AACU],
2015; Presidents Council of Advisors on Science and Technology [PCAST], 2012)at what some might describe as a glacial
pace. When DBER articles are published in high-profile disciplinary research journals there is an opportunity to recruit more
faculty members to reform. Can their impact on the change process be enhanced? Learning theory suggests the answer to this
question is yes.
One model for pedagogical reformhow it occurs, what
promotes it, how barriers can be overcomeis to consider the
process as a learning phenomenon (Mulnix, 2013, 2016). How
people learn is the content. Faculty members and administrators are the learners. Moving from novice to expert in understanding and applying learning theory is the learning goal.
Reformed classroom practice is the outcome at the individual
level; reformed processes and practices are the outcomes at the
institutional level.
FACULTY AS LEARNERS IN THE REFORM PROCESS
Those involved in faculty development have long viewed their
professional roles as facilitating learning, even applying the
term learner to faculty members (e.g., Borko, 2004; McDaniel,
1987). Shulman (1986, p. 4) made the idea of faculty as learners clear for K12 teachers in his seminal article, which examined knowledge growth in teaching. Schulman (1986) argued
that teachers learn not ony about pedagogy (e.g., how to accomplish active learning or group work) but also learn pedagogical
content knowledge (PCK)the knowledge a teacher learns
about teaching in his or her particular discipline (e.g., What are
the likely misconceptions? What are useful metaphors? What
approaches are successful?). Others (e.g., McDaniel, 1987;
Gess-Newsome etal., 2003) have applied Shulmans ideas to
higher education. Major and Palmer (2006) use the lens of PCK
to examine the process of university faculty adopting problem-based learning at the institutional level; their study indicates that professional development activities not only support
learning PCK but can bring about transformative change.
Accordingly, professional developers embed learning theory in their programs (e.g., Sunal etal., 2001; Stewart, 2014).
STEM education researchers also advise use of what is known
about learning to assist faculty members in reform. For
example, Hilborn (2012) advocated for STEM education
CBELife Sciences Education 15:es8, Winter 2016

Faculty as Learners
workshops to incorporate the content of Ambrose etal. (2010)

to better support learning by faculty members. In their report
on change in entrepreneurship education in engineering programs, Giersch and McMartin (2014) similarly argue for faculty development interventions to be informed by learning
theory and principles of instructional design. Kezar (2009)
argues that, for STEM education reform to be successful, faculty must engage in learning that allows sense to be made of
the changes under way.
While those involved in professional development view faculty members as learners, faculty members do not always perceive themselves in this role. Indeed, I have argued elsewhere
(Mulnix, 2016) that helping faculty members see the parallels
between their own learning about learning and their students
learning content is important to transformational pedagogical
change. Table 2 explores this premise of pedagogical reform as
a learning process for faculty by interpreting observations of
STEM education reform from the point of view of faculty members as learners. For instance, the principle that learning is
enhanced by practice with feedback applies to students who
benefit from low-stakes quizzes in ways similar to faculty who
benefit from the kind of formative peer evaluation that occurs
in learning communities.
Table 3 further explores the idea of faculty as learners by
highlighting parallels between faculty members and students as
nave learners. For instance, the fact that faculty members rely
on pedagogies familiar to them from their own prior experiences in education mirrors the fact that students use study strategies that were successful for them in high school. Both populations resist changing their behaviors, even in the face of
evidence that those practices do not optimize learning. Another
example: faculty members, like students, can have incomplete
understandings that are problematic for further learning.
Richardson (1990), Tobias (1992), Michael (2007), Blackie

etal. (2010), Nelson (2010), and Boyd (2014) are among those
who explore nave conceptions held by faculty about learning
and/or reform.
When faculty members are viewed as learners, the impact
of research articles on pedagogical reform can be enhanced.
One way to accomplish this is to explicitly consider research
reports as educational tools and apply what is known about
learning to their content and presentation. Learning science
itself tells us that grounding new information in theory is key
to moving learners along the continuum from novice to
expert. Incoming knowledge that remains disconnected, conceptually unorganized, and isolated from underlying principles is not deeply understood nor easily retrieved (NRC, 2000,
2012a). This knowledge remains inert; it is difficult to use in
new contexts. This means that, for reports of research findings
to have the greatest impact on reform, the learning principles
that might be at work need to be foregrounded and the details
of the experiments and their results need to be better tied to
theory.
Principles are schemes that capture relationships; they elucidate the why and how. They have explanatory power across
multiple contexts: student age, institution, discipline, instructor, class size, and so on. They allow sense to be made of disparate observations such as the facts that reading questions, quizzing before class, identifying muddy points, and reviewing class
notes all enhance learning. That structure determines function
is one example of a broad principle in cell biology; that signal
transduction is typically initiated by a small molecule triggering
protein conformational change is another. While obvious in
definition and example, the principles around which an expert
organizes knowledge are typically unrecognized at a conscious
level (Ambrose etal., 2010). The fluency and automaticity of
TABLE 2. Comparison of student and faculty experiences as learners

Learning principle
Experience of students learning

disciplinary content
Experience of faculty members learning about learning
Learning is dependent on prior Some students arrive at college with low

Faculty members typically receive little/no formal training
experiences and knowledge.
content background and skill development.
before their first teaching jobs.
Learning is a social process.
Students benefit from practices such as think
Faculty members identify lack of collegial and departmental
pairshare, peer instruction, and group work
support as a barrier to change (Brownell and Tanner,
that require students to interact with their
2012).
peers (Kober, 2015).
Teachers identify discussion as important to learning about
learning (King, 2004; Vescio etal., 2008).
Learning environments have
There is a hidden curriculum in traditional
embedded cultural messages.
STEM classrooms that rewards a narrow set
of behaviors, attributes, and prior experiences, namely those associated with lecture.
Making these invisible expectations explicit
can aid students from different cultural
backgrounds (Margolis, 2001).
Learning is maximized by
Performance is improved when students have
practice and abundant,
opportunities to practice recalling and using
timely, and relevant
knowledge in low-stakes evaluations with
feedback.
time to improve performance based on
feedback (NRC, 2012b; Carpenter, 2014).
Significant time on task is

required for deep learning.
Students must spend time truly engaged in the

material to make learning effective.
That the professional identities of faculty members are often

more as scientists and less as teachers is identified as a
barrier change (Brownell and Tanner, 2012).
The particular teaching climate at an institution impacts the
success of professional development interventions
(Condon etal., 2015).
Faculty members benefit from participation in learning
communities that involve shared practice (e.g., Gruppen
etal., 2003).
Gormally etal., 2014 argue that providing faculty with
formative teaching feedback may be the single most
underappreciated factor in enhancing science education
reform efforts (p. 188).
Faculty members identify time constraints as a barrier to
revamping pedagogies (Zakrajsek, 2013).

15:es8, 3
A. B. Mulnix
TABLE 3. Examples of learner statements that illustrate parallels between faculty and students as learners
Examples of learner statements
Faculty
Student
Learners have a knowledge and experience history that is unique. This history can aid learning, or it can hamper it.
I dont understand why todays students need more than a good
lecture and a textbook to learn.
I dont understand why I got a C on the test, I studied my flashcards

and highlighted the reading.
Learners have nave conceptions that limit learning and prevent deep understanding.
Group work results in the weaker students parasitizing the stronger
ones.
Students should readily recall what has been covered in prerequisite
courses.
I teach large classes in auditorium-style classes; active learning just
isnt possible.
Enzymes can make a reaction with a positive free-energy move in the

forward direction.
Plants dont need mitochondria because they have chloroplasts.
Learners have difficulty transferring information learned in one context to another.

I see why providing connections to the lives of students is important,
but there just arent many examples of how molecular biology
relates to real life.
We studied oxidation in organic chemistry, but can you explain again

why it is important in the citric acid cycle?
Learners are more successful when the value of what is being learned is made obvious.
I dont see a need to change my teaching; my teaching evaluations are
quite good.
I didnt really do the reading; the teacher always lectures on whats

in the book.
Statements in the table are not direct quotes but are aggregates that reflect views expressed in workshops with STEM faculty members and teaching-learning experiences
with students.
understanding for an expert are so great that the underlying

structure and process is typically taken for granted unless
explicitly explored (e.g., Feldon etal., 2010).
PRACTICAL APPLICATIONS OF LEARNING PRINCIPLES
In the case of learning, principles offer explanations for why a
particular pedagogical choice is effective. What is it about active
learning or increased course structure or problem solving
during class that might be responsible for an increase in learning? These principles typically trace back to patterns of brain
function and mechanisms of information processing. Below is a
selection of learning principles sufficiently robust to have been
identified in multiple literature reviews and textbooks. These
have been derived from extensive empirical evidence across
multiple disciplines. Two sets of examples are provided to illustrate practical use of the principle. The first is for the context of
student learning. The second set considers how the principles
can be used for faculty development, specifically around discussion of a DBER article.
1. Learners have a knowledge and experience history that is
unique. This history can aid learning, or it can hamper it.
Because prior knowledge is literally written into the physical
connections among neurons (called encoding), unless new
information becomes wired into existing networks it will be
lost. Thus, effective instruction must take into account the
prior knowledge of the learner and appropriately connect to
it (NRC, 2000; Wood, 2009; Ambrose etal., 2010).
For students: An upper-level molecular genetics course might
explore evolution of primate color vision (Jacobs and Nathans,
2009) as a model for consolidating knowledge of loci, alleles,
gene duplication, mutation, and Mendelian inheritance
patterns previously covered in an introductory genetics
course.
15:es8, 4
For faculty: Discussion of a DBER article could begin with

faculty members identifying ways in which their classrooms
already accomplish a suggested intervention such as active
learning.
2. Deep factual knowledge is insufficient for expertise. Expert
knowledge is organized into a highly interconnected conceptual framework that facilitates retrieval and application.
Novices not only have less knowledge, they also lack a
robust framework for organizing information. Without organization and connectedness, knowledge can be present but
not recognized as having relevance when needed (NRC,
2000; Moog etal., 2006; Wood, 2009; Ambrose etal., 2010).
For students: A project that uses information at the National
Center for Biotechnology Information website to investigate
a genetic diseasetracing disruption of the affected gene
from the mutation through the impact on transcription and/
or translation to explain the resulting physiological consequences (Mulnix and Penhale, 1997)can provide an
opportunity for students to clarify and conceptually organize
components of the central dogma of molecular biology.
For faculty: Having identified ways that students engage in
active learning in their classrooms, faculty members could
explain how those activities embody learning principles such
as those listed here and how those activities could be modified to enhance them.
3. Learner motivation matters (Ambrose etal., 2010). While
learners themselves play a significant role in their motivation, so do teachers. Instructors and environments that
prime student motivation aid learning and promote the ability to transfer knowledge to novel contexts (NRC, 2012b).
One means for priming student motivation is to use real-life
examples (Doyle, 2011).

Faculty as Learners
For students: Someone teaching molecular genetics might

ask students to draft and enact a simulated genetic counseling scenario, create posters that market new RNA drugs in a
mock conference for physicians, or prepare evidence about
the role of epigenetics to be given in a mock legislative hearing on funding for preschools.
For faculty: Faculty members could be asked to critique an
assignment description or syllabus and then modify it to better incorporate the ideas from the research article being discussed.
4. Learning is enhanced when novices have opportunities to
explain new information to themselves and to others (Wood,
2009; Ambrose etal., 2010; NRC, 2012b). Elaboration,
whether in written, diagrammatic, or oral form, is a powerful learning tool.
For students: An assignment that asks student pairs to read
a research article on the structure of a protein and then present the findings of that paper in poster format is one means
of engaging students in the process of explanation not only
to themselves but also to their peers (Mulnix, 2003).
For faculty: A discussion of a DBER article could include
pairs of faculty identifying questions that were raised for
them as they read the article. These could then be shared in
the group.
5. Practice with new information is key to retention of learning, and it enhances depth of learning. Practice that is distributed over time has been shown to be more effective
then massed practice such as cramming (Carpenter, 2014;
Dunlosky etal., 2013).
For students: Quizzes, rewriting notes, coverage of topics in
both lecture and laboratory formats, and projects such as
those described in the items above provide opportunities for
practice with feedback (see point 6).
For faculty: Pairs in the discussion group could be asked to
follow up with each other three times over a semester about
how their teaching practices have changed as a result of discussing the research article.
6. Frequent, prompt, and relevant feedback timed to allow for
improvement in performance is key to deep learning (Wood,
2009; Ambrose etal., 2010; NRC, 2012b). This formative
(versus summative) feedback does not need to involve grading and can come from peers, answer keys, supplemental
instructors, and faculty talking aloud through an answer
after students have attempted to answer it.
For students: Giving students opportunities to evaluate a
data set as part of content presentation during class time can
serve as practice with feedback before encountering similar
questions on exams (see the Supplemental Material).
For faculty: Faculty could make peer visits to one anothers
classrooms to observe new techniques being used and then
discuss areas of strength and those needing adjustment.
Alternatively, faculty members might videotape a class and
then do a self-evaluation using materials from Harvards
Best Foot Forward program (http://cepr.harvard.edu/video
-observation-toolkit) as guides. This tool kit is designed for
use in K12, but features are applicable to higher education

as well.
7. Reflection and metacognition assist learners in defining
goals, selecting learning strategies, and monitoring progress
toward those goals. Awareness of ones learning as a process
in progress provides opportunity for greater insight into and
control of that process. (NRC, 2000, 2012b; Wood, 2009;
Ambrose etal., 2010).
For students: Exam wrappers (Lovett, 2013) are one example of strategies to promote reflection and metacognition.
For faculty: Discussion of a DBER article might include
exploring ways in which individual faculty might be resistant to incorporating a change (e.g., giving up control during
an activity, covering less content). Additionally, quiz or
exam questions that measure student learning outcomes
using the alternative technique could be used to assess the
change.
While these principles can be can be articulated in isolation,
they are mutually reinforcing. Connecting new to existing
knowledge can be motivating. Elaboration is a form of practice.
Feedback can support metacognition. Practice can assist in
organizing information according to concepts. Indeed, in discussion of a DBER article by faculty, any one of the suggested
approaches is addressing multiple learning principles.
DBER PUBLICATIONS AND LEARNING PRINCIPLES
INTEGRATED
Table 4 further illustrates how these principles might map onto
the practices empirically demonstrated to enhance learning. For
instance, depending on the specifics of the activity, in-class
problem solving could 1) connect incoming information to students prior knowledge, 2) facilitate organization of knowledge
into a conceptual framework, 3) motivate learners via social
interactions, 4) provide opportunities for elaboration by self
and with peers, 5) offer feedback from peers and/or faculty
members, and/or 6) make the learner aware of learning strategies used by others. Discussion of these or other (see Ambrose
etal., 2010; Mayer and Alexander, 2011; Dunlosky etal., 2013;
Benassi etal., 2014; Brown etal., 2014) principles in the introduction, methods, discussion, and implications sections of
research articles can provide a conceptual framework for readers and assist in their construction of an expert organization of
knowledge.
When authors, for instance, indicate that in-class activities
were designed to promote peerpeer interactions in ways that
elicited feedback for students about their understandings, the
article provides the conceptual structure readers need to make
informed choices in their own classes. Use of clickers and think
pairshare and small-group work to solve problems become rich
interventions rather than activities to include for the sake of
active learning. Explaining that reading questions devised to
help students develop more interconnected knowledge structures
and to connect new to existing knowledge would guide faculty
members as they create and modify supports for their own students. Readers of research articles begin to see that it is not the
particular activity (e.g., clicker questions or thinkpairshare)
that is important but rather how that activity promotes learning.
Understanding the mechanical details of flipped classrooms,

15:es8, 5
A. B. Mulnix
TABLE 4. How learning principles can map onto examples of pedagogical practices
Evidence-based practice
Learning principles that may be at work
Preparation before class:

- providing students with reading questions,
- quizzing before arrival in class
- instruction via video
(e.g., Freeman etal., 2011; Haak etal., 2011; Offerdahlt
and Montplaisir, 2013; Gross etal., 2015)
Creating expert organization of knowledge

Providing cues about the larger concepts and how to hang details on that
framework
Engaging prior knowledge, including misunderstandings
Highlighting common misunderstandings of information
Scaffolding may help students grasp ideas in a logical order or in smaller steps
Active-learning exercises and activities that replace lecture:

- peerpeer teaching
- small group work on conceptual problems
- case studies
- problem solving
- service learning
(e.g., Crouch and Mazur, 2001; Haak etal., 2011;
Freeman etal., 2014; Stockwell etal., 2015)
Prompting self-elaboration and practice

Providing feedback from peers
Engaging prior knowledge (e.g., use of real-world examples)
Motivating students via connection to real-life examples, social interactions,
novelty
Guiding development of expert knowledge organization
Providing feedback via peers
Improving metacognitive skills as students explain their reasoning
Engagement activities that enhance lecture:

- clicker questions
- thinkpairshare
- one-minute papers
- data analysis
- interpretation of diagrams
(e.g., Lovett, 2013; Linton etal., 2014; Smith etal., 2014)
Engaging prior knowledge

Prompting elaboration with self and peers
Providing feedback via discussion and then display of answers
Engaging misconceptions and prior knowledge
active learning, and increased course structure is necessary but

insufficient to allow evaluation, creation, and troubleshooting in
ones own classroom context. It takes an understanding of the
critical attributes of these pedagogical tools to make them effective in other contexts. Richardson (1990, p. 16) argued more
than 20 years ago for education researchers to make their theoretical framework obvious to practitioners so that teachers can
relate the activitys theoretical framework to their own beliefs
and understandings. In this way, teachers have opportunities to
take control of their choices rather than have a given practice
imposed by external forces. Without such empowerment, teachers may become victims of their personal biographies, systemic
political demands, and ecological conditions, rather than making
use of them in developing and sustaining worthwhile and significant change (Richardson, 1990, p. 16).
But dont university faculty have the wisdom and experience
to intuit this conceptual foundation? In fact, there are suggestions in the literature that STEM faculty members do not make
these conceptual leaps. For instance, when Andrews etal.
(2011) studied a broadly representative set of introductory-level biology instructors self-reporting use of active-learning
strategies, little evidence of gains in student learning were documented. This suggests a surface rather than a deep understanding of research findings. Furthermore, instructors modifications of published practices often diminish the effectiveness
of those practices (Dancy and Henderson, 2010). Faculty using
EBP in the classroom have also been shown: 1) to rely on personal over empirical evidence (Cooper, 2014; Andrews and
Lemons, 2015), 2) to have personal beliefs inconsistent with
recommended practice (Borrego etal., 2013), and 3) to misrepresent the degree to which they use reformed practices (EbertMay etal., 2011). The finding that the largest loss of physicists
in reform efforts was at the continuation stage (Table 1; Henderson etal., 2012) also suggests that faculty may lack suffi15:es8, 6
cient depth of understanding of reform practices to personalize

their use.
In a model for reform that views faculty as learners, the difficulties encountered when changing practices built over a
career are not surprising. Difficulty troubleshooting when using
a practice in a new context, returning to prior approaches that
are comfortable, inaccurate estimates of progress in reform, and
questions about the value of the work required are all hurdles
encountered by novices. Even as expert learners in their discipline, most faculty members are novices to the learning sciences. They need the theory behind new practices made evident. The robust literature in the area of job training (e.g.,
McGehee and Thayer, 1961; Baldwin and Ford, 1988; Grossman and Salas, 2011) also indicates that applying learning
principles when designing professional learning/job training is
key to subsequent use of knowledge by employees. In essence,
STEM education reform is a problem of knowledge transfer;
faculty members must usethat is, apply, evaluate, synthesize,
and createtheir knowledge in novel contexts. Such transfer
has been empirically shown to be promoted by a deep understanding of the fundamental principles at work (NRC, 2000,
2012b).
While some DBER articles are thin on theory, there are many
examples that effectively communicate the links among practice and learning science principles. Offerdahlt and Montplaisir
(2013) integrate a robust treatment of formative assessment
into their introduction about how student-generated reading
questions provide a window into student understanding. In
doing so, they not only address feedback but also knowledge
construction, knowledge organization, and metacognition.
Similarly, Gross etal. (2015) use their introductory remarks to
contextualize the roles of student motivation and distributed
practice as underlying principles for improved student performance. Eddy and Hogan (2014) consider increased practice

Faculty as Learners
with material (Freeman etal., 2011) as a factor through which

increased course structure achieves enhanced learning. They
explore time on task, class community, and perceived value of
the course as additional factors contributing to effectiveness.
Linton etal. (2014) explicate why writing may be effective as a
component of active learning, considering knowledge construction and organization, feedback, and metacognition as possible
contributors in their introductory comments. Mack and Towns
(2016) also provide a strong theoretical foundation for their
examination of faculty beliefs about teaching undergraduate
physical chemistry.
My main thesis is that, when theoretical foundations are
overlooked in DBER publications, the impact of the publication
is diminished because readers do not encounter the information that can help them bring new practices to their own classrooms. But also worrisome is the fact that absence of theory
perpetuates misinformation about the learning sciences. It
allows the erroneous belief (e.g., Zhang etal., 2010; Weimer,
2013) that evidence supporting the effectiveness of EBP is relatively new and remains insubstantial to remain unchallenged.
Colleagues and I (Mulnix etal., 2016) have experienced such
misunderstandings in our work with STEM faculty, and hesitancy about quality and quantity of evidence is often implicit in
the sections of papers and reports that continue to address sufficiency of evidence of effectiveness of alternative practices
(e.g., Fairweather, 2008; Appendix F in PCAST, 2012; Mack
and Towns, 2016). While evidence of effectiveness of reformed
practices has not resulted in widespread adoption, removing
doubt in the minds of STEM faculty could reduce a barrier to
reform.
DBER, A MATURING FIELD
Others (Ansari and Coch, 2006; Fairweather, 2008; Hilborn,
2012; DAvanzo 2013; Coppola and Krajcik 2014; Talanquer,
2014) have also noted the need to better integrate DBER with
sister disciplines in the learning sciences. Such integration of
knowledge seems part and parcel of the emergence of learning
science as a discipline. As the field matures, connections among
findings and patterns that are consistent across disciplines
become obvious. The conceptual understanding of the field
deepens.
Indeed, DBER experts themselves can gain insights from
analyzing their own and the STEM communitys reform processes. These reflective and metacognitive insights can then be
used to anticipate the intellectual gaps and challenges that
peers at early stages in reform are likely to encounter. This is
similar to faculty members using disciplinary pedagogical content knowledge to anticipate the misunderstandings that undergraduates are likely to experience. Linking theory and practice
is an area DBER is growing into, with peers pointing out the
need as the field advances. We can learn from this experience:
faculty members are likely to also need support in making these
intellectual connections.
So, how can research publications better contextualize
DBER within learning science theory and empirical findings
from sister disciplines? How can they better support faculty in
the transition to EBP? Attention of authors, reviewers, and
editors to the following aspects of DBER publications are among
the actions that could be taken:
1. Explicitly integrate learning principles into subsections of a

paper, particularly the introduction and conclusion. The
learning principles that could be at work when increased
learning is observed need to be discussed not only to
enhance the quality of the paper but to also better educate
readers.
2. Provide citations to the relevant literature in sister disciplines, not just DBER. The robust evidence that EBP enhances
and deepens learning is summarized in several reviews
(NRC, 2000, 2012a,b) authored by committees of national
experts tasked with integrating and evaluating findings
across the branches of the learning sciences. Available as
free downloadable PDFs from the National Academies Press,
these contain extensive bibliographies. Additional summaries and syntheses with translations to classroom applications are also available (Ambrose etal., 2010; Dunlosky
etal., 2013; Kober, 2015).
3. Report sufficient details that practices can be replicated elsewhere. With the advent of online supplementary materials,
examples of tools such as videos, in-class problems, and
multiple-choice exams can be shared.
4. Supplementary materials should additionally be annotated
so that novice faculty can see the learning principles at work
and have guidelines for implementing them in their own
contexts (e.g., see the Supplemental Material for this article).
Including these items would enrich the context for the
research, provide a more accurate description of the state of the
field, and provide critical resources for novices. The urgency
with which the STEM community needs to accomplish undergraduate educational reform (PCAST, 2012; AACU, 2015)
heightens the need for top-tier journals to model pedagogical
excellence.
CONCLUSION
In summary, educational theory (Ambrose etal., 2010; Sousa,
2010; Mayer and Alexander, 2011; NRC, 2000, 2012b; Fink,
2013) tells us that a deep understanding of content is necessary
for successful transfer of knowledge to novel situations. Such
depth is not only about acquiring information but also about
organizing and connecting it so relevant knowledge is recognized when needed. As learners, faculty members in the process
of reform must not only build their factual knowledge but must
organize that knowledge and experience to allow deep and
transformative learning. An understanding of the learning principles that are the foundation for alternative teaching and
learning strategies can help faculty members develop the ability
to transfer knowledge to their own circumstances. This line of
reasoning suggests that DBER needs to better identify and
explicate learning principles that could account individually or
in combination for improvement on measures of learning. This
is important for assisting novice STEM faculty in their transition
to reformed pedagogical practices.
Because reform efforts are progressing so slowly, publications reporting DEBR results, especially those in high-impact
journals, have a corollary function of professional development.
This function adds an additional responsibility for authors,
reviewers, and editors: to explicitly consider how effective a
paper is as a point of learning for faculty.

15:es8, 7
A. B. Mulnix
DBER experts are pioneers in an emerging field. By explicitly

tying practice to learning principles emerging from sister disciplines, they can greatly help novices in the learning process.
DBER experts have blazed a trail. Creating maps, adding trail
markers, providing signposts, constructing bridges, building
stairs, and adding guardrails will help more people travel the
route successfully.
ACKNOWLEDGMENTS
A.B.M. is supported in part by a grant from the Andrew W.
Mellon Foundation to Franklin and Marshall College.
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15:es8, 9
Supplemental Material
CBELife Sciences Education
S S Mulnix et al.
The three exercises contained in the supplementary materials illustrate how data can be
used in a biology classroom as well as provide examples of how annotations to class
materials can be provided to assist other faculty.

These exercises were created by Amy Mulnix for use in first-year and second-year cell
biology courses.
Non-Covalent Bonding and Geckos

Practice Reading a Research Article
LEARNING GOALS
1. Comprehend the main ideas in a scientific article.
2. Extracting information from highly technical and complex writing.
3. Apply your understanding of non-covalent bonds to larger biological phenomena.
4. Understand how physics and biology can interface.
5. Generate hypotheses.
Article: Autumn, K., Liang, Y.A., Hsleh, T., Zesch, W., Chan, W. P., Kenny, T. W.,
Fearling, R. and Full, R.J. 2000. Adhesive force of a single gecko foot-hair. Nature.
405:681-685.
BACKGROUND
This article discusses the evidence for the mechanism by which a gecko is able to climb
smooth, vertical surfaces. While the article is dense and contains concepts from physics
and engineering, the essential findings are exciting and understandable by introductory
level undergraduate students in biology and chemistry. The article is a wonderful
example of applying interdisciplinary perspectives to a biological problem.
Before beginning to examine the article, generate at least two hypotheses for how a gecko
is able to climb smooth, vertical surfaces.
Because the article is sophisticated, a series of questions are provided here to focus on the
major findings and interpretations relating to the adhesive force of geckos. The process of
gecko detachment from a surface is not considered. Experimental method is not
emphasized but is examined. One goal of this assignment is to learn to read a complicated
article in a selective manner, assimilating information that is familiar, deciding which
information needs to be investigated further, and ignoring information that is not relevant
to the exercise at hand.
A. Read the abstract and answer the following questions.
1. Give a general description of the organization of the hairs (or setae) on a geckos
foot (how many, how long, diameter, shape, etc.). Make a sketch of a seta.
2. What is the definition of a van der Waals attractive force?
3. Before reading any more of the article, write several sentences that explain how
van der Waals forces could account for the adhesion (stickiness) of a geckos foot
to a smooth surface.
4. Examine Figure 1 to obtain visual images of the organization, shape and
dimensions of the setae.
B. Examine Figure 1a, e, and f.
1. There are two types of measurements being made in this paper. One is the force
required to pull when a surface is perpendicular (lift the gecko straight off the
wall). The other is the force that operates in the parallel dimension (the gecko
sliding down the wall, due to gravity for instance).
C. On p. 683 in the first column:
Setal force did, in fact, depend on three-dimensional orientation (spatulae pointing
towards or away from the surface) and the extent to which the seta was preloaded
(pushed into and pulled along the surface) during the initial contact. Contacting the
surface with the seta in a direction other than with spatulae projecting toward the surface
resulted in forces of less than 0.3 microNewtons when the seta was pulled away
perpendicular to the surface. By contrast, when the active spatular region was projecting
toward the surface, force increased enormously. After an initial push toward the surface,
a perpendicular preload, the seta was pulled parallel to the surface. Setal adhesive force
parallel to the surface increased until the seta begin to slide off the edge of the sensor.
This set of sentences is a complicated description of how the seta was applied to the force
sensor. The diagrams at the top of Figure 2 may be helpful in interpreting the information.
The graphical representation in Figure 2a should also be helpful. Put these sentences and
figures into your own description of how the seta is being applied to the sensor and at
what point a strong force is obtained.
D. At the top of the second column on p. 683, the authors begin to make their argument
for van der Waals forces being the source of the ability of geckos to climb smooth
vertical surfaces. Where does this interpretation of results begin (rather than merely
presenting of results)?
E. What are the other two possible explanations for the adhesive forces and what is/are
the negative evidence against these mechanisms?
F. What mechanism does the data in this paper best support for the adhesive force?
G. From what you know of van der Waals forces (particularly that they are additive and
must occur over atomic distances), can you provide an explanation for why an orientation
step (preload) followed by a sliding motion of the foot might provide the greatest
adhesive force in the model begin investigated?
H. Reflect for a moment on how you approached reading this difficult article. What
strategies were helpful? Talk to at least one other person in class and find out what
strategies they used.
I. How does this article connect to things you already know? Can you think of any
practical applications for this work?
ADDITIONAL RESOURCES
1. Video of a man-made material mimic of the gecko foot adhesion process.
http://www.youtube.com/watch?v=HTbrsx1zARs&feature=related
2. This site is at Stanford: http://bdml.stanford.edu/twiki/bin/view/Rise/StickyBot

and contains various information about StickyBot, a robot able to climb surfaces
based on the Geckos mechanism. There is a link to numerous videos showing
3. The web page for the Autumn lab which did the research for this paper is located
at: http://geckolab.lclark.edu/dept/Welcome.html.
http://geckolab.lclark.edu/dept/Welcome.htmlhttp://geckolab.lclark.edu/dept/geck
ostory.html
4. http://bdml.stanford.edu/RiSE/Downloads/Weird_Connections.mov. This does an
excellent job at showing the process of science, including the failures, the testing
of alternative hypotheses and then application of the way the gecko works to the
development of StickyBot. Students could either watch this video either before or
after the assignment.
For Faculty
What does this exercise do?
Note, for an alternative use of this exercise that would span an entire hour or part of a
laboratory session and would emphasize how science is done, see below.
This is indeed a difficult article for introductory level students to read, but the concepts
are relatively simple. One major benefit is that the article emphasizes the importance of
non-covalent transient interactions to organismal behavior and function. The questions
are intended to guide the students through the portions of the article that are most relevant
to a discussion of non-covalent forces. The questions are highly directive given that this
is likely to be among the first time students will attempt to read a research article of this
type beyond the title and/or abstract. This exercise models how an expert scientist might
approach an article of interest that is not directly in her field.
Principles Emphasized
1. Structure determines function at an organismal level.
2. Connections between molecule (even nanoscale) phenomenon and organisms.
3. If the additional resources include an introduction to the StickyBot, then this
exercise also emphasizes the practical applications of science.
Opportunities for Transfer and Connection to Previous Knowledge
Students are asked to use their previous experiences to hypothesize about the forces that
might be responsible for the geckos ability.
Students should connect this article to a previous discussion of van der Waals forces.
A specific question is included to help students connect the ideas in this article to other
things they know or have experienced.
Skill Development
Students are guided through extracting information from a difficult research paper.
They are also asked to examine diagrams, data and statements in the text and attempt to
coordinate these.
Metacognition
The exercise explicitly asks students to consider the approaches they (and a classmate)
took to the exercise that were successful.
How to use this exercise.
Small groups of two - four students could be used in a discussion format. Students could
be assigned the article and given the questions prior to class. If students arrive with some
familiarity of the article, this exercise could take 20 30 minutes, including the
debriefing.
The content debriefing could be done as an oral one with full class participation.
Alternatively, a written set of answers could be distributed at the end of the exercise and
then discussed by the small groups. This approach is preferable to posting the answers for
students to view later as it would allow an in-class discussion on items that were still
unclear.
An oral debriefing on process is highly encouraged, even if it is just the faculty member
reflecting on his or her own approach to the article. Acknowledgement of the difficulty of
the reading is key. Depending on the faculty members goals, this exercise could be used
to emphasize the approach to reading an article more than the content of the article. In
this case, the teacher should collect comments from the students about how the process
went. Some specific questions to ask are:
Write down two or three words that describe how you felt at the outset of the
exercise.
Write down two or three words that describe how you felt while doing the
questions.
Write down two or three words that describe how you are feeling now.
Do you need to understand all of the content of the paper to understand the broad
message?
Did discussing the material with other students help?
Also see below for an alternative to expanding this exercise to illustrate the scientific
process and not just the specific content.
Guidance/Support
Support should be provided for students doing this exercise to minimize their anxiety and
confusion. In many cases, explicit verbal support and acknowledgement that the article
and assignment are difficult but valuable may be necessary. Working in small discussion
groups should alleviate some of the anxiety felt by students. This exercise could also be
given in a recitation or small laboratory setting (see below).
Another way to provide support for students and to keep them moving forward with the
assignment during class is to answer some of the questions posed at intervals during the
discussion. For instance, after giving the students a chance to struggle with the text
associated with Figure 2, a brief explanation to the entire class would be helpful.
Options for grabbing student attention and/or creating multiple sensory cues around this
subject are:
Showing a video section of the StickyBot (see Additional Resources). Illustrate
the 3M Command strips for hanging pictures available at most hardware stores.
The key here is to focus on the surface that attaches to the wall (not the one that
holds the picture). You attach the strip by pushing in a perpendicular direction;
you remove the strip by pulling in a parallel direction. Ask students if it is
possible that a similar mechanism as seen with the geckos is at work.
Alternative Use of this Exercise to Illustrate the Scientific Process

This exercise in combination with the Weird Connections video
http://bdml.stanford.edu/RiSE/Downloads/Weird_Connections.mov would provide an
excellent example of how science works, as well as provide support in understanding the
article. The video explicitly shows development of the idea from an observation in nature
through development of a new technology and then application of that technology for
saving lives. The program acknowledges the difficulties scientists had in figuring out the
mechanism and the scientists themselves talk about how for months they could not figure
out how to get a single seta to stably affix to a surface. It also illustrates the cooperative
nature of the scientific process and how one idea builds on the next.
One way approach for this lesson is to:
1. Give students 10-20 minutes to begin work on the exercise. They should at least
begin the questions in parts A-C. They are likely to find the material difficult and
are likely to be a bit confused.
2. Watch the video through the first approximately 9 minutes (to the point where the
narrator says: And the question is: How?
3. Then let the students return to parts A and B and finish those questions.
4. Give the students another 10-15 minutes to work on parts C-G. Then show the
next approximately 2 minutes in which the notion of van der Waals forces is
discussed. Stop the video at the image of the dog.
5. Ask students to spend 5 minutes discussing part H. Then move them onto
discussing part I.
6. Watch the rest of the video, pointing out how the curling of the toes on the
StickyBot is how the force is released (from the perpendicular rather than the
parallel direction).
7. Finally, pose the question to the students: what do the article and video tell you
about how science is done and how it moves forward?
8. Provide the students with a Command strip for picture hanging and let them
experiment with attaching and detaching it. Asking them to make observations
about both the picture hanging surfaces and the surfaces that stick the strip to the
wall. How are these phenomena like (or not like) the gecko mechanism?
Cystic Fibrosis and Protein Sorting

The majority of mutations associated with cystic fibrosis occur in a protein called the cystic
fibrosis transmembrane conductance regulator (CFTR) protein. This protein is found in the
plasma membrane of many cell types, including those of the lung, sweat glands, and
gastrointestinal tract. The function of the CFTR protein is as a chloride ion (Cl-) transporter.
Many patients with cystic fibrosis have a reduced amount of CFTR in their plasma membrane
and, as a consequence, a decreased Cl- conductance across the membrane. You are researching
the intracellular sorting pathway of one type of mutant CFTR protein in relation to cystic
fibrosis. You perform an analysis to determine the amount of CFTR present in various organelles
from lung cells of normal persons and those of persons suffering from cystic fibrosis. The
following table is a summary of those results. A - means no protein was found in the
organelle, + means that some protein was found in the organelle and +++ means a lot of
protein was found in that location.
NORMAL cells
cytoplasm
ER
cis GOLGI
Trans Golgi network
Plasma membrane
+
+
+++
CYSTIC FIBROSIS
cells
+++
+
+
In one or two sentences, state the location of the protein in cells expressing the normal CFTR
protein.
In one or two sentences, state the location of the protein in cells expressing the CFTR mutant.
From your understanding of protein sorting within the endomembrane pathway, suggest one or
more reason(s) why the mutant protein might have this distribution. In other words, generate a
hypothesis about how the mutation in the CFTR protein of CF patients disrupts its sorting. There
are numerous possible answers to this question; be sure to supply the logic for your suggested
mechanism.
For Faculty
This exercises focuses on protein trafficking in the endomembrane pathway. Students are
presented with a set of idealized data and asked to interpret the results. Their interpretation is
dependent on their understanding of the various factors operating along the endomembrane
pathway. Because the exact dysfunction that leads to accumulation of the mutant CFTR protein
within the ER remains controversial, the question is truly an open-ended one.
How to use this exercise.
This exercise should follow coverage of sorting along the endomembrane pathway, including the
concepts that: 1) protein sorting to the ER is dependent on an ER signal sequence; and 2)
retention/retrieval mechanisms exist for maintaining compartment identity. If faculty elect to
mention/discuss the ER-misfolded protein pathway (also known as ER quality control
checkpoint) by which proteins that do not fold properly are degraded by the cytoplasmic
ubiquitin-proteasome machinery, students will have another aspect of the mutant CFTR to
consider. Faculty may elect to use this exercise as an introduction to a discussion of this latter
pathway.
Students should be asked to work in groups of two to four for this exercise. Although the
answers to the first two questions seem obvious (students are asked simply to describe the
results), some students have difficulty with this step. Faculty may need to provide some
reassurance that there is not a single correct answer to the third question and that if a hypothesis
can be supported by logic, the suggested mechanism by which CFTR accumulates in the nucleus
is a reasonable one.
Guidance/comments for the exercise.
The data has been idealized and summarized. Nevertheless, it is representative of the situation
with the F508 mutant of CFTR: the mutant protein accumulates in the ER whereas a reduced
amount reaches the plasma membrane; little other protein is seen along the secretory pathway.
In addition, the mutant protein that does reach the surface of the cell is rapidly degraded via the
endocytic pathway. The focus of this exercise is on protein sorting through the
ER/Golgi/TGN/plasma membrane. Endocytosis has not been considered for this situation.
Answers
Reasons that the mutant protein might accumulate in the ER include:
1) loss of a signal that directs the protein to the next compartment (Golgi network). Depending
on how in-depth vesicular trafficking has been discussed, students may be able to specifically
suggest that a component of the COP-coated machinery has lost the ability to recognize the
region of CFTR that directs CFTR to the next compartment.
2) the mutation exposes or creates a signal that actively retains the CFTR in the ER
compartment.
3) the mutation disrupts the ability of the protein to assume a final, stable configuration. This
results in the accumulation of the protein in the ER because it does not move forward in the
endomembrane pathway. Eventually, these misfolded proteins are targeted for degradation via
the ER-misfolded protein pathway.
There may be others mechanisms that can be supported by logic.
A recent research article [Tsigelny, I et al., 2005. Identification of molecular determinants that
modulate trafficking of 508 CFTR, the mutant ABC transporter associated with cystic fibrosis.
Cell Biochem. Biophys. 42(1): 41-53.] provides evidence relating to these possibilities. The
abstract of this article is relatively easy to read and presents the possibilities in a succinct
manner. This abstract may be used to supplement the exercise and to provide guidance for or
validation of student thinking. The abstract of this article is a good introduction to reading
scientific literature.
Fates of Pyruvate
LEARNING GOALS
At the end of this exercise you will
1. better understand the pathways for the breakdown of glucose in environments with and
without oxygen.
2. appreciate that cells have options for how they metabolize compounds depending on the
conditions.
3. have practiced generating hypotheses and interpreting data.
PRELIMINARY EXERCISE
What do the following have in common?
Tumor cells
Bacteria at the bottom of a pond
Muscles in the middle of a sprint
Production of beer, wine, cider, sake or soy sauce

Making kimchi, yogurt or sauerkraut
Extended diving in emperor penguins
BACKGROUND
Under aerobic conditions, pyruvate can enter the reactions of pyruvate dehydrogenase
(PDH) and the citric acid cycle. These generate additional ATP (in the form of GTP) and NADH,
as well as FADH2. The NADH and FADH2 ultimately transfer their electrons to oxygen; as a
result of this electron transfer, even more ATP is synthesized. These latter reactions are
absolutely dependent on the presence of oxygen. Thus, the PDH and citric acid cycle reactions
occur only in the presence of oxygen. This set of pathways is often called aerobic respiration.
In contrast, glycolysis can occur under either aerobic or anaerobic conditions. Although
the amount of ATP synthesized is small (only 2), there are numerous anaerobic circumstances in
which glycolysis is the major pathway by which ATP is generated. For instance, in a short,
sudden burst of muscle activity there is a demand for ATP but oxygen is not yet being delivered
via the circulatory system in sufficient quantities to allow aerobic respiration. Glycolysis under
anaerobic conditions produces the ATP necessary to sustain muscle activity for a short period of
time. Another example: yeast living under anaerobic conditions such as those found in the
production of alcohol (beer, wine, liquor).
Glycolysis that occurs under anaerobic conditions is called fermentation. The ATP
produced in this process is used to sustain the cells. Recall, however, that glycolysis also
involves the oxidation of glyceraldehyde 3-phosphate to 3-phosphoglycerate, with the electrons
being transferred to NAD+ to produce NADH. This NADH poses a metabolic challenge to the
cells. In order for fermentation to continue, NAD+ must be regenerated from the NADH. If
NAD+ is not regenerated, then glycolysis will come to a stop. In aerobic metabolism the NAD+
would be regenerated from the electrons being transferred from NADH to oxygen. In anaerobic
conditions a different pathway for transferring the electrons be used to re-oxidize the NADH.
Several metabolic pathways have evolved to accomplish the oxidization of NADH in
anaerobic environments. These are diagrammed on the last page of this exercise.
EXERCISE
This portion of the exercise explicitly works to connect with the chemistry you have been
learning. Examine the attached diagram and complete the following set of questions. The
pathway shown on the far left occurs under aerobic conditions. For this pathway:
1. Is pyruvate oxidized or reduced in this reaction?
2. What is the fate of the acetyl-CoA produced in this reaction?
3. Does this reaction oxidize NADH and regenerate NAD+?
4. How will NADH be oxidized under aerobic conditions?
There are two remaining pathways shown in the center and on the right of the diagram. Look
first at the pathway in the center of the diagram. In this reaction set, which is performed by yeast
under anaerobic conditions, pyruvate is converted to acetaldehyde and then to ethanol.
Examine the first reaction: the conversion of pyruvate to acetaldehyde. Fill in the blank to the
right of the first arrow with the appropriate compound. Examine the reaction that converts
acetaldehyde to ethanol. Fill in the blank with the appropriate compound.
Answer the following:
6. Conversion of pyruvate to acetaldehyde is what general type of reaction?
A. Dehydrogenation
B. Condensation
C. Decarboxylation
D. Hydrolysis
7. Circle the group within acetaldehyde that is altered in the reaction that converts
acetaldehyde to ethanol.
8. Is acetaldehyde OXIDIZED or REDUCED to generate ethanol?
9. Is NAD+ regenerated in this reaction set?
YES
NO
Examine the pathway at the far right of the diagram. This reaction occurs in muscles under
anaerobic conditions. Fill in the blank next to the arrow with the appropriate compound.
10. Circle the group in pyruvate that is altered in the reaction that converts pyruvate to lactate.
11. Is pyruvate OXIDIZED or REDUCED in this reaction?
12. Is NAD+ regenerated by this reaction? YES
NO
13. Is the following statement accurate? Explain.

Under anaerobic conditions, pyruvate can accept electrons, which results in the oxidation of
NADH to regenerate NAD+.
The challenge questions ask you to apply what you have been learning about glycolysis and
fermentation. Each of these questions also asks you to generate a hypothesis and then use data
extracted from research papers to determine whether your hypothesis is supported or not. Note
that the data does not necessarily answer the questions directly; you will have to do some
analysis.
Challenge Question 1
Emperor penguins are among the deepest diving mammals known. They typically dive to 100200 meters; the record depth is over 500 meters. The lengths of most dives are 5-7 minutes, but
some penguins have been observed to stay under for as long as 20 minutes! Remember that
penguins are birds and that birds have lungs not gills. Penguins are not breathing underwater.
Assume you are a researcher studying the physiology of penguin diving. Generate a hypothesis
about which of the two alternative pathways for anaerobic respiration is used by penguins. How
might you test this hypothesis?
The data is from:
Ponganis PJ, Kooyman GL, Starke LN, Kooyman CA, and TG Kooyman.1997. Post-dive blood
lactate concentrations in Emperor penguins, Aptendodytes foresteri. J Exp Biol. 200:1623-1626.
The authors report that the mean value of blood lactate concentration in resting penguins (those
that had not been diving) was 1.70.71mmoll1 (mmol per liter).
The data in the graph below show lactate levels for individual penguins (and two seals indicated
by the WS symbols) beginning at the point that the blood lactate levels peaked after swimming.
This means that the first value shown is the highest lactate concentration reached after a dive for
a given penguin.
Note that the authors of this paper are asking a more sophisticated question about the limits of
diving in the penguins and so they are presenting their data to best show the answers to their
questions. But, you can use the data to decide if your hypothesis is or is not supported.
Does the data presented in the paper support your hypothesis?
Challenge Question 2
As a tumor grows in size, the cells that are most interior in the mass become starved for oxygen
a situation known as hypoxia. This has a limiting effect on the growth of the tumor. One way that
a tumor progresses to a more aggressive state is to produce angiogenic factors. These cause
blood vessels to infiltrate the tumor and connect it to the blood stream. This not only allows for
greater tumor growth (the tumor cells can make more ATP in the presence of oxygen delivered
by the red blood cells), it also provides a route for the tumor cells to migrate around the body (a
process known as metastasis).
Generate a hypothesis about which of the two anaerobic pathways are used by tumor cells when
oxygen is limiting (under hypoxic conditions). How could you test this hypothesis?
Attached is a table taken from S. Walenta, A. Salameh, H. Lyng, J. F. Evensen, M. Mitze, E. K.

Rofstad, and W. Mueller-Klieser. 1997. Correlation of high lactate levels in head and neck
tumors with incidence of metastasis. Am J Pathol. 150(2): 409415.
Just from the title of the article, is your hypothesis supported?
It turns out that advanced tumors produce lactate regardless of whether oxygen is abundant or
not. The reasons behind this are unclear and are still being investigated. This feature is so unique
that lactate levels can be used as a diagnostic feature in staging a tumor. That is to say that
normal tissues do not produce lactate when oxygen is present and if tumor cells are producing
lactate when oxygen is present then the tumor is at an advanced stage. In this article, the authors
make a comparison of tumors of the head and neck that are metastatic with those that are not
metastatic.
Note: The data that is pertinent to this question is in the last column of the table.
Summarize the general results shown in the table. Your answer can be in everyday vocabulary
and should be only a few sentences long.
The authors indicate in their results section that the difference between the metastatic samples
and the nonmetastatic samples is statistically significant. From this data, what can you conclude
about the nature of glycolysis in metastatic tumors as compared to nonmetastatic tumors?
For Faculty
Principles Emphasized
1. In order for catabolic reactions to continue, a supply of NAD+ must be available.
2. Even universal pathways can have different outcomes, depending on the environment in
which the organism evolved.
3. Pathways within organisms are responsive to environmental stimuli.
Connection with Previous Knowledge.
Students are extremely likely to be familiar with the process of fermentation from their life
experiences. If the video is shown (see below), the connection of fermentation and food is even
further emphasized. Students are also likely to recognize other anaerobic conditions (bottom of a
pond and extended diving in mammals). Many students are attracted to penguins and will be
able to relate to the questions relating to their metabolism during dives. Similarly, some students
will have an interest in or experience with tumor biology.
Transfer to New Contexts.
Depending on content covered previously in the course, students will have encountered the ideas
of oxidation and reduction. This is an opportunity to use those concepts in a new context. This
exercise, if used between discussing glycolysis and the citric acid cycle, will also introduce
students to decarboxylation reactions.
Skills Development
Students have the chance to practice interpreting diagrams of chemical reactions. If the option is
taken (see below), students are also asked to apply their knowledge to a real-life scenario,
generate a hypothesis and determine if their hypothesis is supported by a real data set.
How to use this exercise
Students can be given the first portion of the exercise either in-class or as a homework
assignment. If given as an in-class assignment, 10-12 minutes should be allowed for students to
complete the exercise working alone. If an emphasis has been placed on recognizing the
chemistry of glycolysis and the citric acid cycle, students should be able to complete this
assignment on their own. Indeed, this assignment can be used as a formative assessment tool; if
students have trouble completing the exercise, they need to continue working with the concepts.
It is important to debrief students at the end of this first section before doing the challenge
exercises (if that option is taken). This immediate feedback is necessary to correct any
misunderstandings before moving to the next portion.
The second portion of the exercise uses the challenge questions. The intent is to allow students:
1) to connect with previous knowledge (about penguins and tumors respectively); and to practice
hypothesis generation, interpretation of data and drawing conclusions. These could be given as
homework or done in class.
Guidance/comments for the exercise

Many but not all introductory cellular/molecular texts include a discussion of fermentation. The
introductory comments for this exercise should be sufficient to allow completion of this exercise
even if the text does not cover these concepts.
This first portion of the exercise is intended to be used when an emphasis has been placed on
understanding and recognizing the chemistry of glycolysis and the citric acid cycle (e.g.,
decarboxylation, phosphorylation, oxidation, etc.). Students that have not previously been asked
to recognize a decarboxylation or oxidation reaction are likely to have difficulty with this
assignment.
The second portion of the exercise could be used on its own if the proper content has been
presented previously. Students could complete this in class or as homework. This may be an
appropriate approach in an upper level course.
During the time that students are working, faculty and/or teaching assistants should circulate. A
major goal is to see where students are getting stuck. If there is a common misunderstanding, this
can then be addressed immediately. Circulating in this manner also helps faculty better
understand the students strategies for completing the assignment. When helping students, the
goal should be to direct their attention to a feature (such as how many carbons are in each
compound or what are the differences between the reactant and product?) rather than providing
an answer directly.
The entire exercise could be used as an alternative to a lecture on this material. An outline for
how that might be done is:
Ask students what the list of items has in common.
Watch the video on fermentation in food (link is below).
Complete the portion of the worksheet on the possible fates for pyruvate. Faculty may
wish to model how to approach the exercise by talking through their approach of the
pyruvate to acetyl-CoA series. Students would then work the two pathways to the left.
Model for students how to generate a hypothesis and look for evidence by asking them
which of the two anaerobic processes they believe is happening in the baking of bread
and the fermentation of ginger ale (examples given in video). Then ask for one piece of
data from the video that supports their hypothesis (answer = giving off CO2 gas causing
the bread to rise and the pressure to increase in the bottle of ginger ale). You could also
extend this to beer and wine making with the piece of evidence being that alcoholic
fermentation is happening because the product is ethanol.
Let students work the challenge questions. Students could chose which topic they want to
work with (both the tumor and penguin examples deal with production of lactate).
When the students have finished working their exercise they should turn to their neighbor
and discuss their work (it wont matter if their neighbor did the same or different topic).
Debrief both the penguin and tumor scenarios with the entire class. Place an emphasis on
data interpretation. Note on the graph provided in the penguin example that the WP
points are not for Emperor penguins. Be sure to identify the axes on the graph and discuss
the quick decrease in lactate in the first 10 minute followed by a leveling off in the
remaining time. You could also discuss the individual variability in the data.
End class with a brief discussion of the facts that production of lactate is only a
temporary solution to anaerobic conditions in mammals since its production disrupts the
acid/base balance in tissues. You may want to incorporate the following facts: 1) humans
too produce lactate during exercise; the lactate is sent to the blood stream and this must
be metabolized by the liver (this is the Cori cycle); 2) part of the way that tumors are
located is by providing a radioactive sugar compound and asking where the radioactivity
shows up in a PET scan tumors are one of the major tissues using the sugar and so they
light up readily.
Options for grabbing student attention and/or providing multiple sensory cues.
This video explains fermentation in the context of food. The first half of the video is sufficient
for the explanation. The second half shows how to make ginger ale. It is a bit kitschy but fun.
http://www.youtube.com/watch?v=tNqfPsVAdYk. Of course you can serve ginger ale as
students come to class.
The introduction to this second video provides some facts on Emperor penguin diving (the
remainder of the video discusses the speakers unusual path to research in biology).
http://www.youtube.com/watch?v=VPUetsnURVk A few pictures of diving penguins are at the
Cool Antartica website.
http://www.coolantarctica.com/Antarctica%20fact%20file/wildlife/Emperor_penguin_pictures.ht
m.
Answers.
The following modifications should be made to the diagram. A CO2 should be placed in the
blank to the right of the arrow from pyruvate to acetylaldehyde. NADH to NAD+ should be
placed in the blanks to the right of the arrow from acetylaldehyde to ethanol. An NADH to
NAD+ should also be place to the right of the arrow from pyruvate to lactate.
1. Pyruvate dehydrogenase
2. Oxidized
3. Goes to citric acid cycle
4. No
5. NADH will be oxidized by passing its electrons into the electron transfer chain and finally
onto oxygen.
6. C. Decarboxylation
7. The ketone group is reduced to a hydroxyl group
8. Reduced
9. Yes
10. The ketone group is reduced to form a hydroxyl

11. Reduced
12. Yes
13. NAD+ is regenerated under aerobic conditions by passing electrons in to the electron
transport chain and then to oxygen.
14. Ethanol, carbon dioxide, lactate
15. The statement is accurate but incomplete. Pyruvate directly accepts electrons (is reduced)
only in the case of its conversion to lactate.
Penguin Scenario
Possible hypothesis: Penguins will convert pyruvate to lactate while they are diving and are not
breathing. Testing the hypothesis would involve measuring lactate in their bodies before and
after diving.
The data in the statement from the authors and the graph indicate that lactate levels increase in
the penguins during their dives. This suggests that glycolysis is following the lactate pathway to
regenerate NAD+ when oxygen is not available (has been used up).
Tumor Scenario
Possible hypothesis: Tumors produce lactate under anaerobic conditions. This could be tested by
measuring the lactate present in metastatic tumor (or in the blood) and comparing it with the
lactate levels in persons nonmetastatic tumors.
It is clear from the last column in the table that lactate is produced by tumors; and the amount
produced by metastatic tumors is higher than that produced by nonmetastatic tumors.

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ESSAY

STEM Faculty as Learners in Pedagogical

CONTRIBUTIONS OF DISCIPLINE-BASED EDUCATION RESEARCH

Deborah Allen, Monitoring Editor

Supplemental Material can be found at:

Tried vs. not

High user vs.

low-risk, private environments and to troubleshoot new

Stains (2015, p. 23) report that DBER articles are largely

CBELife Sciences Education 15:es8, Winter 2016

workshops to incorporate the content of Ambrose etal. (2010)

Richardson (1990), Tobias (1992), Michael (2007), Blackie

TABLE 2. Comparison of student and faculty experiences as learners

Experience of students learning

Experience of faculty members learning about learning

Learning is dependent on prior Some students arrive at college with low

Significant time on task is

Students must spend time truly engaged in the

That the professional identities of faculty members are often

CBELife Sciences Education 15:es8, Winter 2016

I dont understand why I got a C on the test, I studied my flashcards

Enzymes can make a reaction with a positive free-energy move in the

Learners have difficulty transferring information learned in one context to another.

We studied oxidation in organic chemistry, but can you explain again

I didnt really do the reading; the teacher always lectures on whats

understanding for an expert are so great that the underlying

For faculty: Discussion of a DBER article could begin with

CBELife Sciences Education 15:es8, Winter 2016

For students: Someone teaching molecular genetics might

use in K12, but features are applicable to higher education

CBELife Sciences Education 15:es8, Winter 2016

Learning principles that may be at work

Preparation before class:

Creating expert organization of knowledge

Active-learning exercises and activities that replace lecture:

Prompting self-elaboration and practice

Engagement activities that enhance lecture:

Engaging prior knowledge

active learning, and increased course structure is necessary but

cient depth of understanding of reform practices to personalize

CBELife Sciences Education 15:es8, Winter 2016

with material (Freeman etal., 2011) as a factor through which

1. Explicitly integrate learning principles into subsections of a

CBELife Sciences Education 15:es8, Winter 2016

DBER experts are pioneers in an emerging field. By explicitly

Dunlosky J, Rawson KA, Marsh EJ, Nathan MJ, Willingham DT (2013).

Cooper MM (2014). Evidence-based reform of teaching and learning. Anal

Haak DC, HilleRisLambers J, Pitre E, Freeman S (2011). Increased structure

Coppola BP, Krajcik JS (2014). Discipline-centered post-secondary science

CBELife Sciences Education 15:es8, Winter 2016

Kezar A (2009). Synthesis of scholarship on change in higher education.

Offerdahlt EG, Montplaisir L (2013). Student-generated reading questions:

Presidents Council of Advisors on Science and Technology (2012). Engage

Pusateri TP (2014). How SoTL can contribute to faculty development and

Richardson V (1990). Significant and worthwhile change in teaching practice.

McGehee W, Thayer PW (1961). Training in Business and Industry, Hoboken,

Michael J (2007). Faculty perceptions about barriers to active learning. Coll

Talanquer V (2014). DBER and STEM education reform: are we up to the

Tobias S (1992). Revitalizing Undergraduate Science: Why Some Things Work

Mulnix A (2013). Communicating a new model: learner-centered strategies

Vescio V, Ross D, Adams A (2008). A review of research on the impact of

Weimer M (2013). Learner-Centered Teaching, San Francisco, CA: JosseyBass.

Mulnix AB (2016). What my cadaver dog taught me about teaching and