FALL 2024 VOLUME 70 NUMBER 3

PLANT SCIENCE 

BULLETIN

A PUBLICATION OF THE BOTANICAL SOCIETY OF AMERICA

In This Issue

The Development of BSA’s 

Comprehensive AI Policy for 

Its Academic Journals....p. 205

New Editors-in-Chief Named: 

Sean Graham (

AJB) & 

Carolina Siniscalchi (

PSB) .... p. 197

Insights from a Fulbright U.S. 

Scholar Alumni Ambassador 

by Nishanta Rajakaruna .... p. 277

A Pair of SciArt articles 

focused on paleobotany! .... p. 259

Congressional Visits Day Report

by Jenna Miladin 

& Cael Dant... p. 286

A Collection of Articles by 

Charles E. Bessey Teaching 

Award Winners ... p. 216

                                    Fall 2024 Volume 70 Number 3

FROM the EDITOR

Sincerely,

Greetings, 

Putting together this issue of Plant Science Bulletin has been bittersweet for me, as it is my 

last issue as editor-in-chief. Serving as PSB editor has been one of the highlights of my career 

and I speak more about this in a short question-and-answer segment on page 199. I am very 

proud of this issue, as it exemplifies my favorite kind of PSB issue by including a little bit of 

everything. We have timely articles that focus on pressing issues in botany, including one by 

Theresa Culley and colleagues that addresses the use of AI in BSA publications and one by 

Caroline Bose that discusses why and how botany can become more inclusive and accessible. 

You will also find reflections by the two winners of the 2024 BSA Public Policy Award who 

attended the AIBS Communication Boot Camp and Congressional Visits Day in D.C. to 

promote science and botany to legislators. 

The highlight of this issue, for me, is a special feature on education in which several of our 

recent Charles E. Bessey Award winners share teaching philosophies and strategies. I was 

thrilled with the diversity of articles that I received in response to my invitations. I am 

also happy to include two final articles on the theme of science and art. Both articles look 

specifically at the role of art in paleobotany.  

Thanks to you all for being readers of Plant Science Bulletin. I hope you enjoy this issue!

PSB 70 (3) 2024

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195

TABLE OF CONTENTS

SOCIETY NEWS

Changes in Editors-in-Chief for Two BSA Publications..................................................................197

Ten Years of 

Plant Science Bulletin

: An Exit Interview with Editor-in-Chief 

MackenzieTaylor.......................................................................................................................................199

An Exit Interview with

American Journal of Botany

 Editor-in-Chief Pamela Diggle

         After a Decade of Service ....................................................................................................................202

The Development of BSA’s Comprehensive AI Policy for Its Academic Journals........205

SPECIAL SECTION

HONORING THE TRADITION OF BOTANY EDUCATION IN THE 

PLANT SCIENCE 

BULLETIN: 

A COLLECTION OF ARTICLES BY CHARLES E. BESSEY TEACHING 

AWARD WINNERS...................................................................................................................................................216

Four Things I Learned from 30 Years of Teaching

          (that you probably already know) (by Cynthia Jones) ........................................................218

Universal Design for Learning Botany (by J. Phil Gibson)............................................................223

Field-based courses still matter, but not like they used to 

          (by Christopher T.  Martine)...............................................................................................................227

Neo-natural history:  careful observation and co-discovery in 

          teaching botany.(By Joan Edwards) .............................................................................................230

Using Inquiry as a Tool to Help Students Develop a more Sophisticated Understanding 

           of Frequently Misunderstood Concepts. (by Marshall Sundberg)...............................235

Don’t forget our roots: learning with drawing.  (By Stefanie M. Ickert-Bond 

           and Brett C. Couch)....................................................................................................................... 242 

The two rules of great teaching: present with enthusiasm and make your 

           students do the work (By Bruce Kirchoff)  ........................................................................... 248

The Evolution of an Educator – (By Suzanne Koptur).................................................................. 252

ART IN THE BOTANICAL SCIENCES: PAST, PRESENT, AND FUTURE..........................259

         Illustrating Cretaceous Park: First Steps Toward a Botanical Field Guide for the 

               Hell Creek Formation (By Kirk R. Johnson and Marjorie Leggitt)................................260

Reconstructing the Botanical Past: Art and Paleobotany (By Edward J. Spagnuolo, 

          L. Alejandro Giraldo, Mario Coiro, and Susannah Lydon) ................................................264

SPECIAL FEATURES

The Fulbright U.S. Scholar Program: Insights from a Fulbright U.S. Scholar 

          Alumni Ambassador ..............................................................................................................................277

Twelve Pounds of Duct Tape and No Manual: Shifting Mindsets Around Disability in   

          Botany............................................................................................................................................................286

Report from 2024 Congressional Visits Day ........................................................................................290

PSB 70 (3) 2024

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196

TABLE OF CONTENTS

 MEMBERSHIP NEWS

BSA Virtual Symposium on Climate Change: .....................................................................................290

Support Graduate Students with Year-End Donations to the GSRA Fund

          Donate today!.............................................................................................................................................290

Help Us Reach Our Goal of 100 Gift Memberships by December 31! .................................291

Three-Year Memberships—Stay Connected at a Discount!........................................................291

Botany360 Updates............................................................................................................................................291

BSA Sponsorship Opportunities.................................................................................................................292

BSA Student Chapters.....................................................................................................................................292

BSA Spotlight Series.........................................................................................................................................292

FROM THE 

PSB ARCHIVES.............................................................................................................294

SCIENCE EDUCATION

Student Perceptions of Scientists: Preliminary Results from PlantingScience 

         F2 Research Project...................................................................................................................................295

Master Plant Science Team Applications Open for Spring and Fall 2025..........................300

Huge PlantingScience Fall Session Wrapping Up............................................................................300

State-by-State Resource Update: List of States/Territories Still Needed............................300

Nominations for 2025 Bessey Award.......................................................................................................301

STUDENT NEWS

Botany 2024 Recap............................................................................................................................................302

Grant Opportunities............................................................................................................................................302

Grad School Advice ...........................................................................................................................................302

Papers to Read for Future Leaders ...........................................................................................................303

ANNOUNCEMENTS

Art, Ecology, and the Resilience of a Maine Island: The Monhegan Wildlands................301

IN MEMORIAM

Pieter Baas(1944–2024) ................................................................................................................................305

Dr. Elisabeth Zindler-Frank ...........................................................................................................................307

BOOK REVIEWS......................................................................................................................................308

PSB 70 (3) 2024

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197

The Botanical Society of America is thrilled to 

announce that Dr. Sean Graham (University of 

British Columbia) will serve as the new Editor-in-

Chief for the American Journal of Botany (AJB) 

and Dr. Carolina Siniscalchi (Mississippi State 

University) will serve as the new Editor-in-Chief 

of the Plant Science Bulletin (PSB) beginning in 

January 2025. 

Both Drs. Graham and Siniscalchi bring to 

their new roles impressive credentials and 

strong commitments to Society publications. In 

concordance with the strategic goals of the BSA, 

Changes in Editors-in-Chief for 

Two BSA Publications

DR. SEAN GRAHAM APPOINTED NEW EDITOR-IN-CHIEF OF THE 

AMERICAN JOURNAL OF BOTANY AND DR. CAROLINA SINISCALCHI 

APPOINTED NEW EDITOR-IN-CHIEF OF THE 

PLANT SCIENCE BULLETIN

they both are committed to diversity, equity, and 

inclusion as an essential practice in all aspects of 

science.
Dr. Graham is a Professor in the Department 

of Botany at the University of British Columbia, 

Vancouver, Canada, who has wide-ranging research 

interests in plant systematics and evolution, and in 

particular characterizing plant biodiversity from 

phylogenetic and phylogenomic perspectives. His 

interests have ranged from addressing challenging 

higher-order relationships—both across and 

within the major lineages of land plants—to more 

focused systematic studies of closely related taxa. 

SOCIETY NEWS

DR. SEAN GRAHAM

DR. CAROLINA SINISCALCHI

PSB 70 (3) 2024

198

He has studied the molecular evolution of plant 

genes and genomes, and the evolution of plant 

sexual systems. He has strong ongoing research 

interests in monocots and mycoheterotrophic 

plants.
In addition to a full career as a professor and recent 

Head of the Botany Department at UBC (2016–

2021), Dr. Graham has served BSA Publications 

for many years, including as an AJB Associate 

Editor (2008–2017 and 2019–2024), and as a guest 

co-editor on two AJB special issues (“Exploring the 

Potential of Angiosperms353, a Universal Toolkit 

for Flowering Plant Phylogenomics” in 2023; the 

Charles Darwin Bicentennial in 2009).  He has 

also played a publications-related leadership role, 

as he was elected for two successive terms as the 

BSA Director-at-large, Publications. In this role he 

helped lead the transition of AJB and Applications 

in Plant Sciences from self-publishing to 

partnering with the commercial publisher Wiley. 

As a Director, he was also a BSA board member, 

and he advocated to the Board for the creation 

of the AJB Synthesis Prize for early-career 

researchers (ECRs). He has also served in multiple 

additional official and unofficial service roles, 

including on the BSA publications committee and 

the publication ethics subcommittee. He regularly 

assists the editorial team with analysis of the annual 

Journal Impact Factor and has strongly promoted 

the need to increase the number of review articles 

as a key tool to improve our impact more broadly. 

This insight helped lead to the creation of an AJB 

“Reviews Editor” role at the journal, and was part 

of the motivation to establish the AJB Synthesis 

Prize..
According to Dr. Graham, “I believe strongly 

in society-run scientific journals, which are 

motivated by science over profit. I therefore 

regularly publish some of my best research in AJB. 

I would like to find new ways to encourage others 

to do so, too.” 

Dr. Graham will begin his five-year term on 

January 1, 2025. He replaces the remarkable 

current Editor-in-Chief, Dr. Pamela Diggle, 

whose second five-year term concludes December 

31, 2024. [See her outgoing thoughts elsewhere in 

this issue of the PSB.]

Dr. Siniscalchi is an Assistant Professor and Data 

Science Coordinator in the University Libraries 

at Mississippi State University. Her main areas of 

botanical research interest are the macroevolution 

of the nitrogen-fixation symbiosis in flowering 

plants and the systematics and evolution of 

the sunflower family. She also has expertise 

in data science, bioinformatics, and research 

data management. Her strong background in 

systematics research and current position in 

library science are a unique combination that will 

bring new ideas and directions to the PSB. 

Dr. Siniscalchi received her bachelor’s, master’s, 

and doctoral degrees from the Universidade 

de São Paulo, Brazil. She has been a member 

of the BSA since 2017, when she first moved to 

the United States, and has attended five Botany 

meetings since then. She was a member of the 

APPS Reviewing Board from 2020 to 2022, served 

on the BSA International Affairs Committee 

(2019–2021), and is currently the Secretary/

Treasurer for the Southeastern Section. 

Dr. Siniscalchi’s vision for the PSB is that it will 

reflect the wide array of interests and diversity 

of BSA’s membership. “I want BSA members to 

see the bulletin as not only the place where they 

receive information from the society, but also as 

the place where they can talk about themes that 

are not strictly scientific but that are inherently 

part of being a botanist (and I use botanist here in 

the widest sense: not only as academics, but every 

person that has plants as the center focus of their 

work or hobby),” she says. 

Dr. Siniscalchi will begin her five-year term on 

January 1, 2025. She replaces the amazing current 

Editor-in-Chief, Dr. Mackenzie Taylor, whose 

second five-year term concludes December 31, 

2024. [See her outgoing thoughts elsewhere in this 

issue of the PSB.]

PSB 70 (3) 2024

199

Ten Years of 

Plant Science Bulletin

: 

An Exit Interview with 

Editor-in-Chief Mackenzie Taylor

What first drew you to take on the role of editor-

in-chief of the Plant Science Bulletin?

I was interested in serving as the editor-in-chief 

of PSB because I believe strongly in its role as a 

resource for the botanical community. I have 

always loved the variety of articles in the PSB and 

the fact that it celebrates the achievements of BSA 

members. I think, at its best, it builds community 

within the BSA and provides a place for important 

discussions to occur outside of annual meetings. 

Additionally, I wanted to provide a positive 

experience for others who wished to publish 

in  PSB, especially for people rather new to 

publishing. My first publication was in PSB

(Johnson et al., 2004), and I valued the experience 

of getting to work with collaborators and publish 

an article as an undergraduate. Marsh Sundberg, 

who was PSB editor at the time, made this a very 

positive experience and I hoped to pass this along 

to others. 

What were your goals as editor-in-chief?

During my time as editor, I have had three 

primary goals for the PSB. The first has been to 

provide a platform for members and friends of the 

BSA to share ideas and knowledge in the realms 

of education, public policy, public outreach, and 

history. I consider the PSB to be the publication of 

record for the BSA in matters outside of scientific 

research. I believe that its pages should provide a 

snapshot of the environment in which botanical 

research and education is taking place, both for 

contemporary readers and for posterity. 

The  PSB team and I have accomplished this by 

inviting many of the people who have given 

addresses to the Society or led workshops, either 

at the Botany meetings or through the Botany360 

program, to prepare written articles so that they 

might reach a broader audience. Some of our most 

thought-provoking pieces have come from these 

contributions. I have also encouraged our Public 

Policy Committee to keep the Society updated 

on matters such as funding for plant science 

research and relevan

t bills that come before 

Congress. When I started as editor, I felt strongly 

that the PSB could play a larger role in promoting 

and facilitating science advocacy. I think we made 

gains in this area. 

PSB 70 (3) 2024

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My second goal was to provide resources for the 

botanical community, especially as they related 

to goal number one. During my 10 years, PSB has 

published articles with practical strategies and tips 

for preparing articles for publication, avoiding 

predatory publishing, submitting successful NSF 

grants, applying for Fulbright awards, conducting 

field work, improving scientific presentation 

skills, and moderating scientific sessions at 

conferences, among many other topics. PSB

authors have contributed to the debate on issues 

such as plant awareness disparity and whether 

standardized tests should be used in admissions. 

We have continued to publish articles that 

present strategies for teaching in the classroom 

and laboratory, as well as for public outreach. 

Further, we created a section just for students. 

The student representatives share information and 

resources for student members and highlight the 

accomplishments of those members. I hope that 

PSB readers have found these articles to be useful.

They continue to be available in the PSB archives. 

My third goal was to elevate as many individual 

voices in the PSB as possible and provide a 

platform for many perspectives. There is always 

room for improvement in this area, but we have 

made a significant effort to engage with the broad 

botany community. For example, our recent 

special issues on Art and Botany included an open 

call for articles; the response was tremendous, 

including from authors who had never published 

in PSB before. In another example, we asked the 

larger community, including on social media, 

for articles about dealing with the pandemic that 

stimulated many thoughtful responses.

How has the direction of the PSB evolved over 

the past 10 years?

Over the last 70 years, PSB has been continually 

evolving to fit the needs of the BSA. During some 

periods it has included more articles and essays 

and in others, it has been more of a newsletter used 

for disseminating news and announcements. Over 

the last 10 years, we have continued a trend to 

reduce the emphasis on news and announcements, 

mostly because these are more easily and quickly 

disseminated via the email newsletters. In turn, 

I have made a deliberate effort to increase the 

number of peer-reviewed articles in each issue. 

My goal was always one or two articles per issue 

and most of the time we accomplished this. I also 

wanted to diversify the type of articles published 

in  PSB so that we were serving as much of the 

botanical community as possible.  

We decided when I became editor that we would 

continue to emphasize the print version as most 

of our readers indicated that they preferred 

that format. Near the start of my first term, we 

revamped the look of the PSB and created a new 

logo that I absolutely love. Ten years on, much has 

changed in the publishing landscape and the new 

editorial team will have to decide if it is time to 

transition to online-only publication or if there are 

new and better ways to reach readers. Whatever it 

looks like in the future, I am hopeful that PSB will 

only grow in value to BSA members. 

What do you consider your most rewarding 

accomplishments in your role with the PSB?

There are many things I’m proud of regarding my 

role as PSB editor. One of the most rewarding to 

me personally was the series of issues that came 

out in 2020–2021. These were very volatile times, 

with universities and businesses shut down due to 

the COVID-19 pandemic and the United States 

in the middle of significant political upheaval. I 

conceptualized and coordinated both the Summer 

and Fall 2020 issues from my dining room table 

because Creighton’s campus was closed. Despite 

this, I believe these issues are some of the most 

important in PSB’s history. We provided tips for 

educators and researchers who were working with 

reduced resources and attempted to provide a 

record of these times for future reference through 

special features (Taylor, 2020; Min et al., 2020; 

Gaynor and Valdez, 2020). We also did our best 

to lift up the voices of people who had timely and 

meaningful ideas to share about inclusion and 

equity in botany (e.g., Dewsbury, 2020; Leonard, 

2020; Asai, 2021) and have made featuring these 

perspectives an ongoing priority. I am also very 

proud of the Art in the Botanical Sciences special 

PSB 70 (3) 2024

201

issues that were published in Fall 2023 and Spring 

2024, although the hardest work was done by the 

guest editors for these issues and the authors. 

These were very well received and demonstrate the 

unique ability PSB has to cross disciplinary lines. 

What has been the best part of serving as PSB

editor?

Serving as the editor of Plant Science Bulletin

has truly been a highlight of my career. I have 

found great joy in thinking about what topics 

members of the Society would be interested in 

and then working with Richard Hund to figure 

out how to best feature that in PSB. The best 

part has been getting to interact with people I 

might not otherwise have had a reason to get to 

know, including our wonderful authors, section 

contributors, and article reviewers, as well as our 

book reviewers and the publishers who provide 

books for review.

Do you have any last thoughts?

It takes a team to create the PSB, so I want 

to thank everyone who has contributed to the 

Bulletin during the last 10 years, whether as an 

author, contributor, reviewer, or book reviewer. 

I especially want to recognize all the student 

representatives and policy committee chairs who 

have prepared sections for each issue as a part 

of their service in that role, as well as Catrina 

Adams and Jennifer Hartley for preparing our 

regular feature on Science Education. Truly, Plant 

Science Bulletin does not exist without those of 

you who contribute your time and energy to PSB. 

Thank you to Amy McPherson and to the BSA 

Publications Committee for helping to develop 

procedures and sharing thoughtful ideas. Special 

thanks to Johanne Stogran for compiling and 

formatting each issue. She does an incredible job 

making the PSB look fantastic in print and ISSU 

format. Finally, it has been an absolute joy to work 

with Richard Hund, PSB managing editor. I know 

that the PSB issue will be in the capable hands of 

Carolina Siniscalchi, and I am excited to see what 

she does to move the Bulletin forward. 

REFERENCES

Asai, D. 2021. The little red hen and culture change. 

Plant Science Bulletin 67 (3): 174-180.  https://bot-

any.org/userdata/IssueArchive/issues/originalfile/

PSB_67__3__2021.pdf
Dewsbury, B. 2020. What have we learned? Lessons 

and strategies from the chaos. Plant Science Bulle-

tin 66 (3): 198-205.  https://botany.org/userdata/Is-

sueArchive/issues/originalfile/PSB_2020_66_3.pdf
Gaynor, S., and I. Valdes. 2020. Shifting gears: field-

work, benchwork, and greenhouse studies during 

COVID-19.  Plant Science Bulletin 66 (3): 236-242.   

https://botany.org/userdata/IssueArchive/issues/origi-

nalfile/PSB_2020_66_3.pdf
Johnson, E. E., M. Taylor, R. Lopez-Smith, 

and D. Morningstar. 2004. What Works for

Me: Undergraduate Perspectives on Professional De-

velopment.  Plant Science Bulletin 50 (4): 102-104.  

https://botany.org/psbarchive/issue/2004-v50-no-4.

html
Leonard, B. G.  2020. Diversity and inclusion in the 

sciences: relationships and reciprocity.  Plant Science 

Bulletin 66 (3): 191-197.  https://botany.org/userdata/

IssueArchive/issues/originalfile/PSB_2020_66_3.pdf
Min, Y., S. Gaynor, and I. Valdes. 2020. Student ex-

periences during the COVID-19 lockdown. Special 

Feature Plant Science Bulletin 66 (2): 129-134. https://

botany.org/userdata/IssueArchive/issues/originalfile/

PSB_2020_66_2.pdf
Taylor, M. 2020. COVID-19 and you: checking in with 

12 BSA members during a global pandemic. Special 

Feature. Plant Science Bulletin 66 (2): 93-110.  https://

botany.org/userdata/IssueArchive/issues/originalfile/

PSB_2020_66_2.pdf

Art in the Botanical Sciences Special Issues

https://botany.org/userdata/IssueArchive/issues/origi-

nalfile/PSB%2070(1)%2020242.pdf

https://botany.org/userdata/IssueArchive/issues/origi-

nalfile/WebPSB_69_3_2023.pdf

PSB 70 (3) 2024

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What first drew you to take on the role of 

AJB Editor-in-Chief?

One (perhaps glib) answer is that Carol Goodwillie 

asked me to!!  She was the BSA’s Director-at-Large 

for Publications and chairing the search committee. 

She was passing through my neighborhood and 

dropped by to discuss the possibility.  I have vivid 

(very positive) memories of the occasion.  We 

went for a long walk and talked at great length 

about the possibilities for the position.  The other, 

more heartfelt, answer is that I had always been 

(and continue to be!) a strong supporter of the 

BSA and had participated in many aspects of its 

governance; I saw the editorship as an important 

opportunity to continue serving the society and 

botany.  The American Journal of Botany also is 

of great significance to me personally.  AJB was 

the first journal I subscribed to as a beginning 

graduate student, and I read the articles avidly. 

The growing row of issues, then bound in bright 

yellow card stock, arranged on my bookshelf, 

gave me a sense of belonging and professionalism.  

My first research paper was published in AJB.  I 

also knew that AJB has been equally important 

in the careers of botanists across the country and 

internationally.  So, as soon as Carol raised the 

possibility,  I got very excited by the prospect and 

immediately began to consider what I might (aim 

to) do as Editor-in-Chief.

What were your goals as Editor-in-Chief?

I looked back at some of the documents I 

submitted with my application for the position and 

this sentence stood out: “The primary challenge 

An Exit Interview with

American Journal of Botany

Editor-in-Chief Pamela Diggle 

After a Decade of Service

faced by the AMERICAN JOURNAL OF BOTANY

is the same challenge faced by the publications 

of all scientific societies: How will the Journal 

maintain relevance in this rapidly evolving world 

of diverse outlets for dissemination of science?” 

This is as true today as it was 10 years ago, and I 

continue to keep my focus on this challenge.  One 

of my goals as incoming EiC was to increase the 

breadth of research areas included in the journal 

and to expand the geographic, institutional, and 

demographic diversity of authors and editors.  To 

this end, I aimed to increase all aspects of diversity 

among the board of Associate Editors.  AJB

currently has 64 Associate Editors, 49% of whom 

are women, and who are in institutions from 

Argentina (1), Austria (1), China (2), Colombia 

(1), Denmark (1), France (2), Germany (4), India 

(1), Israel (1), Korea (1), Mexico (3), Netherlands 

(1), New Zealand (1), North America (39), South 

Africa (1), 

Spain (2), and Sweden (1).  

The 

diversity of authors is more difficult to gauge, but 

PSB 70 (3) 2024

203

we will begin to track self-reported demographic 

data next year and will be able to keep tabs on 

how we’re doing.  I also aimed to “have the pulse 

of the readership”; to understand how readers and 

authors discovered articles, and what new and 

exciting research was on the horizon.  I began 

to implement this immediately by holding a 

series of listening sessions (online and at Botany 

conferences) with botanists from a broad range of 

disciplines and career stages to understand how 

best to serve our community.  These listening 

sessions have now been formalized as the ECAB 

(Early Career Advisory Board), which consists of 

advanced graduate students and post-docs who 

provide input and suggestions through regular 

meetings.  I also wanted to bring new readers and 

authors to the journal by introducing a “News and 

Views” section in each issue of the journal that 

includes non-technical summaries of research 

papers (“Highlights”), brief essays on new areas of 

research (“On The Nature of Things,” now a regular 

feature of most issues), and a diversity of opinion 

pieces and commentaries.  Also, as a result of a plan 

hatched during a 2-day strategy retreat, AJB now 

features regular review articles.  I thank incoming 

EiC Sean Graham, who was at that meeting, for 

presenting a compelling argument for a reviews 

section in AJB.  And I am so very grateful to 

Jannice Friedman for taking on the enormous task 

of getting this feature off the ground successfully 

and serving as Reviews Editor for over two years, 

and to the current Reviews Editor, Kasey Barton, 

for carrying on this important work with grace 

and enthusiasm.

How has the direction of AJB evolved over 

the past 10 years?

Both  AJB and the scholarly publishing industry 

in general have undergone tremendous change 

over the last decade.  The year I started, 2015, 

marked a full century of AJB publication and 

in all of that time, it had been self-published. 

Library subscriptions largely supported the 

journal and, critically, other activities of the 

BSA.  In acknowledgment of changing financial 

models and challenges of competition among 

scientific journals for diminishing resources 

in library budgets, AJB, in 2017, entered into a 

partnership with Wiley and is now in a second 

five-year contract. We joined with Wiley at a time 

when they had a strong stable of Society journal 

partners, and we benefit from their scholarly 

publishing expertise and economies of scale. 

With Wiley we have been better able to adapt to 

the strong push in STEM toward Open Access, 

which offers great advantages but also tremendous 

financial challenges—for both authors and 

Societies. Major changes in AJB’s distribution 

have also occurred.  In 2015, AJB was provided 

to members electronically and/or, by request, as 

a hard copy of each issue. Printing of the journal 

was discontinued in 2019, and now all access is 

electronic. The ability to promote and share links 

to articles to a vast international community of 

botanical enthusiasts was greatly expanded as 

social media exploded over the past decade. As the 

social media landscape has grown more complex, 

AJB, along with the BSA, is emphasizing more 

diverse, and less divisive, platforms. One thing 

that hasn’t changed is our careful copy editing, and 

the care and attention to detail that the AJB staff 

bring to each article and to our authors. 
I want to emphasize, that although publishing has 

undergone dramatic transformations and many 

new features have been added to the Journal, AJB

is a Botanical Society of America publication, 

and as EiC, I have kept the mission to serve the 

society and to publish “peer-reviewed, innovative, 

significant research of interest to a wide audience 

of scientists in all areas of plant biology” in mind 

with every decision that we’ve made.

What do you consider your most rewarding 

accomplishments in your role with 

AJB

?

I would like to highlight my efforts to increase 

AJB’s inclusivity.  As noted above, one of my goals 

as EiC has been to increase the diversity (in all of 

its multiple meanings) of authors, readers, and 

editors. To further this goal, all Special Themed 

Issues now include an open call for proposals 

for articles to be included in the issue. Early-

PSB 70 (3) 2024

204

career and other underrepresented groups of 

authors are especially encouraged to participate.  

And, last year we ran an open call for Associate 

Editors.  This initial call drew a gratifying array 

of applicants from across the globe who were 

interested in serving botanical research generally 

and the journal specifically.  The demographics 

of the BSA and of science and society at large are 

changing rapidly and that should be reflected in 

our journal.  Moreover, encompassing a broad 

and diverse range of perspectives and approaches 

is imperative for addressing the pressing issues of 

global climate change.

What has been the best part of serving as 

AJB editor?

Serving as the Editor-in-Chief of AJB has been one 

of the most gratifying and rewarding experiences 

of my career.  AJB is so much more than a journal.  

It is a community of exceedingly talented people 

working selflessly to advance botanical sciences 

and to support botanical scientists.  We all 

know that, despite our best efforts as authors, it 

is the rare paper that is not improved during the 

peer-review process.   I have had the pleasure of 

watching this “evolutionary process” as reviewers 

and Associate Editors take the time from their 

already over-scheduled days to carefully read 

and comment—some even going so far as to 

suggest new analyses, and provide code and all!  

The result is inevitably a stronger/clearer paper 

with greater impact. We receive many notes from 

authors about the positive experience they had 

at AJB.  I’m grateful for the generous work of all 

the many people involved, past and present, in 

the American Journal of Botany.  AJB’s Associate 

Editors continually amaze me.  They bring such 

knowledge and insight to the papers they handle, 

and each of them is dedicated to the success of 

the journal.  And then, there are the multitude 

of reviewers who cannot be thanked enough for 

their contributions.  A special thanks goes to the 

amazing AJB Managing Editor, Amy McPherson, 

who has very much been my partner over the 

past decade (and the leader in understanding 

the rapid changes to the publishing industry).  

It has also been my pleasure to work with the 

equally amazing Production Editor, Richard 

Hund

, 

who handles all the “behind the scenes” 

complexities that turn your manuscripts into 

published papers.  (And who gleefully smuggled 

chips and beer into the Botany meeting venue for 

our first several information-gathering sessions.)  

Talented Content Editor Staci Nole-Wilson (and 

past Content Editors Sophia Balcomb and Marian 

Chau), among other things, skillfully ensures that 

your papers have all of the vital sections and are 

ready publish.  And, most especially, I thank all 

of you who have contributed to the success of the 

journal by sending your research papers to the 

journal!

PSB 70 (3) 2024

205

In today’s society, artificial intelligence (AI) is 

rapidly advancing and expanding through all 

aspects of our lives. The release of ChatGPT in 

November of 2022 made AI accessible to anyone 

with a computer and an internet connection. After 

the explosion of interest and activity that followed, 

AI now has the potential to radically change 

our world as we know it. According to a recent 

Oxford University Press poll (Anderson, 2024), 

researchers across scientific disciplines today are 

increasingly using AI tools, but also have extensive 

misgivings about AI technology. For example, 76% 

of researchers globally currently use some form 

of AI in their research (e.g., chatbot, machine 

translations, AI-powered search engines and 

research tools), but only 8% trust the AI companies 

not to use their own data without permission, and 

25% are concerned about AI reducing the need 

for critical thinking skills in science (Anderson, 

2024). Most recently, publishers Taylor & Francis 

and Wiley agreed to sell access to academic content 

The Development of BSA’s 

Comprehensive AI Policy for Its 

Academic Journals

By Theresa M. Culley

1,11*

, Irene Cobo-Simón

2*

, Robert L. Baker

3

, Aaron S. David

4

, Matthew A. 

Gitzendanner

5

, Matthew S. Olson

6

, Tilottama Roy

7

, William N. Weaver

8

, Pamela Diggle

9

, and 

Briana Gross

10

1

 Department of Biological Sciences, University of Cincinnati, OH

2 

Institute of Forest Sciences (ICIFOR-INIA, CSIC), Madrid (Spain); Associate Editor of APPS

3

 National Park Service, Fort Collins, CO; Associate Editor of APPS

4 Archbold Biological Station, Venus, FL; Associate Editor of AJB

5 

Department of Biology, University of Florida, FL

6

 Department of Biological Sciences, Texas Tech University, TX

7

 Department of Biology, Missouri Western State University, MO

8

 Department of Ecology and Evolutionary Biology, University of Michigan, MI

9

 Department of Ecology & Evolutionary Biology, University of Connecticut, CT; Editor-in-Chief of AJB

10

 Swenson College of Engineering and Science, University of Minnesota-Duluth, MN; Editor-in-Chief of APPS

11

Corresponding author: theresa.culley@uc.edu

*Joint first authors

to certain tech companies for training AI models, 

causing concern among the scientific community. 

AI itself is a broad term that refers generally to 

non-human (machine) intelligence (De Waard, 

2023), but AI can be adapted and used for specific 

purposes (Zhou, 2023). Underlying many AI tools 

are large language models (LLMs), which are 

trained on large amounts of existing text data or 

visual and sound recordings to decipher written 

human language and create media. LLMs are most 

useful for translation, summarizing existing text, 

and generating requested content such as Q&A. 

Generative AI tools such as ChatGPT use these 

LLMs with additional training to then create 

original content such as text, images, code, and 

even videos or music. AI can also be used in a 

process known as “inference” to draw conclusions 

from new data without depending upon only past 

examples.

PSB 70 (3) 2024

206

Generative AI tools are already impacting multiple 

fields of scientific research and the publication 

of scientific articles. Generative AI tools include 

a wide variety of technologies, such as natural 

language processing (NPL), which underlies 

generative pre-trained transformer (GPT) 

models, and image generation and editing. These 

tools can be used in writing to suggest text, correct 

grammar or spelling, or match a particular style 

of a scientific journal. AI tools are also extremely 

useful for data analysis; they can process large 

amounts of data with accuracy and speed, and 

identify patterns and information difficult to 

detect with traditional methods. AI can be used 

to generate code, automate repetitive tasks, and 

simulate experimental conditions. When used in 

these ways, AI has the exciting potential to propel 

science forward in ways we can only imagine 

today; however, its use also raises important 

ethical and practical considerations. Present-day 

AI-generated content can sometimes include 

incorrect, out-of-date, or nonexistent citations, 

or contain repetitive or inappropriate language, 

reflecting the biases/inaccuracies of the data on 

which the tools have been trained. AI tools can 

be used to manipulate images and may plagiarize 

existing text, but this technology can also be used to 

detect such actions with ever-increasing accuracy. 

For example, publishers such as Elsevier, Springer, 

and Wiley now use their own in-house AI tools 

to check for AI usage in submitted manuscripts to 

ensure integrity of their publications.

Recognizing the necessity of addressing the 

use of AI in the publication process, the 

Botanical Society of America (BSA) formed 

an ad hoc committee in fall 2023 to develop a 

policy regarding use of AI in its publications 

(American Journal of Botany, Applications in Plant 

Sciences,  and  Plant Science Bulletin). Committee 

members consisted of researchers selected from 

a special call for participants, BSA editorial staff 

(managing editors, production staff, associate/

reviewing editors, and editors-in-chief), and the 

BSA Director-at-Large for Publications. This 

committee was charged to discuss generative 

AI tools as they apply to publishing and to then 

develop guidelines, policies, and best practices for 

authors, reviewers, and editors of BSA journals. 

The committee specifically focused on the 

following three categories: 

1. Defining how authors may or may not use AI 

when writing text, including how to properly 

acknowledge AI tools (if allowed in any cir-

cumstance)

2. Describing how AI tools can be used for gen-

erating code as a potentially acceptable use

3. Deciding how reviewers may or may not use 

AI in their reviews

The committee met several times during the 

following months as individual workgroups 

focused on drafting sample language for each 

point above, and then as the full group to fine-

tune the language. This AI policy established 

guidelines to promote responsible and ethical 

use of AI in scientific publications—aiming to 

harness the potential of AI while safeguarding the 

integrity of scientific research. The AI policy was 

then added to the Author Guidelines for all BSA 

journals and released publicly in spring 2024, with 

required disclosure of AI use on the author and 

reviewer submission forms. As AI continues to 

evolve, ongoing dialogue and adaptation of these 

policies will be crucial to ensuring that the BSA 

community remains at the forefront of innovation 

and ethical practice.

The purpose of this article is to describe the key 

points considered by our ad hoc committee during 

our discussions, namely: (1) how other journals 

and publishers have addressed AI to date, (2) 

current opportunities and challenges of AI tools, 

and (3) a summary of our committee discussion 

that resulted in the final BSA AI policy.

PSB 70 (3) 2024

207

 CURRENT STATUS OF AI 

IN PUBLISHING

Here we review as of April 2024 the current 

guidelines and policies of the top six academic 

publishers, as identified by Scholarly Publishers 

Indicators 2022 (https://spi.csic.es/), on the use of 

AI generated content (AIGC):

• Cambridge University Press

https://authorservices.wiley.com/ethics-

guidelines/index.html

• Elsevier

https://www.elsevier.com/about/policies-and-

standards/the-use-of-generative-ai-and-ai-

assisted-technologies-in-writing-for-elsevier

• Oxford University Press  

https://academic.oup.com/pages/authoring/

books/author-use-of-artificial-intelligence

• Taylor & Francis  

https://asset.routledge.com/

rt-files/AUTHOR/Guidelines/

Manuscript+preparation+guide.pdf

• Springer

https://www.springer.com/gp/editorial-poli-

cies/artificial-intelligence--ai-/25428500

• Wiley-Blackwell

https://authorservices.wiley.com/ethics-

guidelines/index.html

All publishers consider the use of AI an ethical 

issue. For example, Oxford University Press states, 

“AI must be consistent with the Press’s mission 

and the values inherent in our publishing, with 

all that this entails in terms of quality, integrity, 

and trust.” All six publishers agree that AI is a 

tool that simulates human intelligence, but is not 

an intelligent entity in itself. Consequently, none 

of the publishers allow a statement of authorship 

by any AI-based tool (such as ChatGPT) in 

scientific articles. This is consistent with the 2023 

statement from the Committee On Publication 

Ethics (COPE; https://publicationethics.org/cope-

position-statements/ai-author), which states that 

AI tools cannot perform the role of an author of 

a work, nor therefore, appear in the list of authors 

of a work. As non-legal entities, AI tools cannot 

take responsibility for the ethical and legal aspects 

of the submitted work. Furthermore, Wiley and 

Elsevier point out the difference between the use 

of AI to make original intellectual contributions 

(without human direction)—which is not 

allowed—versus assistance in the preparation 

of scientific articles—which is allowed. Both 

publishers also point out the need for the authors 

to supervise the content generated by the AI tools. 

All publishers (except Oxford) state that authors 

are ultimately responsible for their manuscript 

content regardless of whether AI was used.

All publishers also agree that the use of AI to 

generate content must be transparent and correctly 

referenced, as required with any other tool. Any 

use of AI must be disclosed in the cover letter 

to the editor upon manuscript submission and/

or in the Methods or Acknowledgments section 

of a manuscript. This is also consistent with 

COPE’s position statement on AI tools. Elsevier, 

Cambridge, and Taylor & Francis all state that the 

use of AI tools must comply with editorial policies 

on authorship and principles of publishing ethics 

(also mentioned in COPE’s position statement). 

Cambridge also emphasizes its anti-plagiarism 

policy, pointing out that any content generated 

by other authors and coming from AI-based tools 

must be cited and referenced in an appropriate 

and transparent manner.

There is a lack of consensus regarding the 

generation or modification of images through AI 

tools. Elsevier and Springer consider AI-generated 

figures separately from the generation of other 

types of content such as text, and prohibit it, with 

few exceptions. While Elsevier does not provide 

any explanation for this policy, Springer supports 

their policy by stating that legal issues relating to 

AI-generated images and videos remain broadly 

unresolved; consequently, Springer is unable to 

permit its use for publication. In contrast, Oxford 

evaluates AI-generated images in a similar way to 

the generation of other types of content (e.g., text, 

PSB 70 (3) 2024

208

code), allowing it as long as it meets the criteria of 

transparency and is cited correctly. The remaining 

publishers do not consider the use of AI to 

generate and/or modify images separately in their 

Author Instructions; therefore, it is understood 

that they consider images along with generation of 

content in general. This is also in line with COPE’s 

position statement on authorship and AI tools, 

which considers AI-generated images similarly 

to other AI-generated content (text, graphical 

elements, data collection, and analysis) and allows 

it as long as authors are transparent in disclosing 

within the article how the AI tool was used and 

which tool was used. Authors are also considered 

fully responsible for any AI-generated content, 

including all of its ethical aspects. 

Several publishers have also developed policies 

concerning the use of AI in the review process. 

Springer stresses transparency in the use of AI 

tools during the peer-review process, requiring 

reviewers to declare any use of AI in their peer-

review report. Springer notes that this technology 

still has considerable limitations (e.g., as 

described below, such as outdated information). 

Furthermore, Springer also explicitly prohibits 

reviewers from uploading any manuscript content 

into generative AI tools because manuscript text 

may contain sensitive or proprietary information. 

Both Elsevier and Springer note the rapid 

advancement of AI tools and therefore the need 

to regularly review their AI-related policies and 

guidelines.

More recently, publishers Taylor & Francis and 

Wiley separately gave licensing rights to AI 

companies for their repository of past publications 

(Dutton, 2024); Oxford University Press and 

Cambridge University Press are now forming 

partnerships as well (Wood, 2024). Taylor & 

Francis’ $10 million deal with Microsoft is expected 

to assist their development of Copilot, Microsoft’s 

AI assistant. Wiley’s partnership with at least two 

undisclosed companies was reportedly worth $23 

million and $21 million; in return, Wiley provides 

access to its published material to train LLMs by 

using book content and small pieces of individual 

articles, and to make a narrow range of articles 

specific to a topic available for use in inference. At 

this point, it is unknown whether authors will even 

know if their publication has been used. Except for 

a few publishers, authors are not able to opt-out 

of having their material used in this way, which 

has created much consternation for many authors 

(Authors Guild, 2024). In the case of Wiley, the 

company has established guiding principles for AI 

technology and partnerships (https://www.wiley.

com/en-us/terms-of-use/ai-principles). 

OPPORTUNITIES OF AI 

TECHNOLOGY

Artificial intelligence and LLMs offer many new 

and exciting opportunities for researchers not 

only to enhance their science, but also to promote 

communication through the publication process 

(Buriak et al., 2023). One of the most common 

uses of AI by authors is as a “personal copy editor” 

to improve the quality and clarity of the language 

in their manuscript, polishing text created by the 

author. When used properly, these tools are not 

dissimilar to automatic spell checkers and grammar 

checkers. Even Microsoft Editor is now promoted 

as an AI-powered service. The popular Grammarly 

tool also boasts of an AI communication assistant 

to help authors pinpoint areas of weakness, such 

as typos, missing punctuation, or commonly 

confused words. The premium version of 

Grammarly is advertised as using AI to adjust the 

tone, rewrite full sentences, and generate text for 

over 1000 different AI prompts in manuscripts and 

even email. Other AI-based editing and rewriting 

tools include Wordtune (for rewriting, shortening, 

or expanding content), WordRake (which edits for 

brevity or simplicity), Writefull (helping to write 

and paraphrase scientific text), and LanguageTool 

(a grammar checker specialized for multilingual 

writers). More grammar checker and rewriting 

tools will undoubtedly be released in the future, 

especially as generative AI and the machine 

learning on which it relies continue to improve.

PSB 70 (3) 2024

209

Such personal copy editors powered by AI may 

be especially helpful for multilingual authors 

for whom English might not be their primary 

language, particularly when submitting to an 

English-only journal. Some authors already 

upload their own text into ChatGPT and then 

review the rewording, grammar, or punctuation 

suggestions to enhance the clarity of their papers. 

ChatGPT can be used for any language within 

its repertoire, which now includes at least 50 

languages, with more being added to make this 

tool increasingly accessible and useful. Currently, 

some AI-suggested text may still be scientifically 

nonsensical or inaccurate, so a careful eye is 

required before accepting and incorporating any 

recommendations (see below). However, with 

continued training, future renditions of AI tools 

will likely overcome these problems. 

AI tools can also be used by researchers to explore 

the literature when first embarking on a new 

topic, and to identify suitable references for their 

manuscript. When asked to provide peer-reviewed 

papers on a specific topic, ChatGPT provides a short 

list of usually five papers, but can be prompted to 

retrieve more. As with all AI-generated results, the 

papers may or may not relate to the topic and need 

to be reviewed further. Recent papers are usually 

excluded from the list, as dates of retrieved papers 

reflect when the AI was initially trained. For 

example, ChatGPT-4 Turbo released in November 

2023 can only identify literature published up to 

April 2023. Other AI-powered platforms such 

as scholarcy (https://www.scholarcy.com/) help 

authors quickly summarize and organize articles 

applicable to their own research, increasing the 

efficiency with which researchers can search the 

literature.  

As more authors use generative AI for polishing 

existing text, there are multiple downstream 

benefits. First, the overall written quality of 

manuscripts submitted to journals may increase, 

making it easier for editors to ascertain if 

a manuscript is appropriate for the journal 

and should be sent out for external review. A 

well-written manuscript is more likely to be 

perceived favorably by reviewers, who can 

focus on the scientific content rather than 

distractions of misspellings, grammatical errors, 

confusing sentence construction, and general 

disorganization. Such a manuscript will also 

reduce the amount of copy editing and time 

required for conversion into a publication-quality 

article, increasing the efficiency of the publication 

process.

For several years, publishers and editorial staff 

have been using their own AI tools to detect 

plagiarism and image manipulation, and to find 

appropriate reviewers for submitted manuscripts. 

BSA journals commonly use CrossRef's Similarity 

Check to review manuscripts for potential 

plagiarism. AI-powered platforms such as 

Proofig or imagetwin can also be used by editors 

and publishers to detect image manipulation. 

Publishers are now piloting AI to detect submitted 

papers generated from “papermills”—groups of 

individuals or an organization generating similar 

papers and submitting them fraudulently to 

multiple journals for financial gain. Editors can use 

AI to analyze a submitted manuscript's relevance 

to a journal, verify the identity of an author, and 

detect irregular publishing patterns by authors 

that may indicate fraud (e.g., a mathematician 

submitting papers to a medical journal). In a 

time where there are increasing numbers of 

predatory journals (Culley, 2018), AI can also 

be used to check the quality of references cited 

within an article. Publishers are also beginning to 

use AI tools to flag machine-generated content, 

especially when text may be translated into one 

language and then converted back in an effort 

to avoid detection (such as “big data” in English 

translated to “data grande” in Spanish and back to 

“greater data”). In a time when finding appropriate 

reviewers willing to read a submission is critical 

to the peer-review process, publishers are now 

using AI tools to locate suitable reviewers or to 

identify conflicts of interest (e.g., a proposed 

reviewer recently co-authoring a paper with the 

author) instead of a handling editor spending 

their own time to track down this information. In 

summary, incorporating AI tools to assist editors 

PSB 70 (3) 2024

210

and publishers can greatly decrease the amount of 

time spent per manuscript, while enhancing the 

quality of the review and publication process. 

Finally, when properly trained, AI technology 

can also be used to effectively conduct science. 

For example, AI-based models can be used to 

synthesize vast quantities of data that would 

otherwise require multiple people and many hours 

of labor. Such synthesis also minimizes the chance 

of mistakes being made and enhances consistency 

of any particular process. The power of AI can also 

be harnessed to identify patterns and relationships 

within large data sets that would otherwise be 

difficult and time consuming to detect. For 

example, LLMs can now be used to interpret text 

in digitized images of herbarium specimen labels 

(Weaver and Smith, 2023; Weaver et al., 2023). 

Another example is the revolutionary and recently 

developed AI program AlphaFold 3, which is 

able to predict the structure and interactions of 

proteins with other molecules such as DNA and 

RNA with unprecedented precision and accuracy 

(Abramson et al., 2024). AI can also be used as an 

additional overlay to identify any information that 

otherwise would regularly go undetected. Finally, 

AI can check code or even generate code within an 

experiment that would take a human many hours 

to create. In summary, the advantages of using 

AI within the scientific process itself are many, 

provided of course that all results are supervised 

and checked by the researcher themselves.

CHALLENGES OF AI 

TECHNOLOGY

While AI poses exciting and innovative 

opportunities, it is not without serious concerns 

and challenges in the publication process, 

particularly when used incorrectly. Many of these 

concerns can be avoided by treating AI as a tool 

to assist human decisions and by recognizing the 

inherent limitations of AI, most of which reflect 

the underlying machine-learning and training 

technology. 

 On the most basic level, AI technology can be prone 

to inherent errors such as incorrect, nonsensical, 

or blatantly false output (Davis, 2023). Citations 

may be incorrect, incomplete, or outdated because 

the AI tool is limited by its most recent training 

date. AI can also be weak at judging whether an 

unusual outcome is “spurious, anomalous or 

groundbreaking” (Buriak et al., 2023). Even the 

ability to detect a typical outcome will depend 

solely on the data provided to the tool during its 

training—hence the strength of any current AI 

tool will always be temporally and contextually 

limited. AI-generated tools are also known for 

sometimes creating shallow and superficial text 

with a superfluous tone. There are now detectors 

that can be used to identify such AI-generated text, 

such as Turnitin, TraceGPT, Hive, and GPTZero, 

but their effectiveness, accuracy, and cost can vary 

(Walters, 2023). In addition, inadvertent errors 

could occur if generative AI incorporates phrases 

that are not in the author’s native language that 

may have an alternative meaning in another 

language that is not understood by the author (e.g., 

“background research” vs. “doing research in the 

background”). Finally, while AI can be effective 

at summarizing past studies (assuming it is able 

to detect all relevant content), the technology at 

the current time is still unable to look forward in 

time and provide a critical assessment of a topic 

and articulate next steps. These types of errors 

are especially concerning if readers assume AI-

generated text is of human origin (Buriak et al., 

2023). Such inaccurate information would also be 

very worrisome if it escapes detection by reviewers 

and is then published in a peer-reviewed journal, 

earning a scientific stamp of approval. In short, 

current AI technology is limited because it lacks 

human intuition and the ability to detect nuances 

and to conclusively project into the future. 

Another major concern with the use of AI 

technology in the publication process involves 

confidentiality. When reviewers are asked to read 

a submission for a peer-reviewed journal, they 

must agree to confidentiality and not share the 

author’s work or ideas. However, confidentiality 

PSB 70 (3) 2024

211

could be violated if a reviewer uses generative AI 

to compose their written review by uploading part 

or all of the submitted manuscript into an AI tool. 

Although this is now starting to change, some 

popular AI tools may still incorporate text that 

has been entered into the search window in the 

subsequent training of its tool or technology, such 

that the same text or idea could potentially be 

suggested by the tool to another user in response 

to a related query. A potential solution is the use 

of private generative AI tools within individual 

laboratories in which training data are kept in-

house; however, even a private generative AI tool 

may suffer from many of the same challenges 

outlined above. Using a private tool to generate 

a brief summary of the manuscript, as typically 

presented at the top of a formal review, could be 

helpful though, provided the platform is used with 

human oversight.  

AI tools may also express inherent biases based 

on the algorithm and training data used to create 

the tool. Such bias can be sexist, racist, or even 

political, depending on what content was used 

in the initial training. For example, ChatGPT 

replicated gender bias when asked to construct 

recommendation letters for males (which used 

nouns such as “expert” and “integrity” and 

adjectives like “respectful” and “reputable”) and 

females (emphasizing “beauty” or “delight” and 

who were “stunning” and “emotional”) (Wan et al., 

2023). In another example where ChatGPT was 

asked to create a crime drama, researchers used 

four-word prompts, only one of which changed 

(either “black” or “white”), to explore ChatGPT’s 

potential implicit bias (Piers, 2024). Motoki et 

al. (2023) also found that ChatGPT exhibits left-

leaning political tendencies, such as towards 

Democrats in the United States, the Workers’ 

Party in Brazil, and the Labour Party in the United 

Kingdom. The reason for these biases is that many 

LLMs use data from the internet for their training, 

which largely reflects historical stereotypes and 

perspectives already present online. Thus, if left 

unchecked, the use of ChatGPT and other LLMs 

could inadvertently amplify existing and historical 

information on the internet and social media. 

 DEVELOPMENT OF THE BSA 

POLICY ON AI TECHNOLOGY 

Our ad hoc committee met several times in 2023 

and 2024 to discuss the ethical use of AI in the 

publishing process. We examined every aspect 

of the development of a research project: initial 

conceptualization, data collection, integration 

and analysis, interpretation and presentation 

of data, and writing the manuscript. Going 

into these discussions, many of our committee 

members were initially skeptical of using AI in the 

publication process due to its inherent limitations 

(see above) and the possibility of authors using it 

unscrupulously to fabricate text. In fact, several 

of us started the conversation thinking about 

excluding all elements of AI from the publication 

process but, as explained below, eventually we 

changed our minds. Ultimately, we agreed that 

there was no part of the scientific process for which 

AI should be banned because it has the potential 

to help in every aspect, if used appropriately. 

We recognized that there is no AI tool that is 

inherently beneficial or detrimental; it depends on 

how a given tool is used and the extent to which 

the user is aware of each tool’s limitations. AI has 

the potential to make research more thorough 

by uncovering additional information beyond an 

author’s immediate knowledge. Thus, we agreed 

that the development of guidelines for authors 

and reviewers for the publication process is key 

to taking advantage of this novel and promising 

technology, while avoiding its potential drawbacks.

We also recognized that the AI field is rapidly 

advancing with constantly evolving tools such 

that what we perceive today as cutting-edge may 

quickly become routine in the months and years to 

come. The AI of tomorrow will likely be different 

from the AI of today because machine learning 

algorithms and technology are rapidly improving. 

Consequently, our committee understood that 

any AI publication policy developed now will 

need to be revisited and modified in the future as 

AI technology changes. 

PSB 70 (3) 2024

212

CONSIDERATIONS FOR 

AUTHOR GUIDELINES

We all agreed at the onset that AI cannot be an 

author because a non-human entity cannot take 

responsibility for a paper. There must be human 

oversight of any AI assistance; it is imperative for 

authors to take full responsibility for any inclusion 

of AI-generated material in their research studies 

and manuscripts. Just as before AI was available, 

we trust authors to adhere to ethical standards 

while conducting their studies and writing their 

manuscripts. However, we also recognize that 

guidance and specific policy are necessary to 

prevent any intentional or inadvertent violations 

within the new AI landscape. Just as it is critical to 

specify when AI is not allowed, it is also important 

to spell out any approved uses of AI tools. We 

largely agreed with what publishers have already 

been doing: AI tools can enhance the quality of 

a manuscript in terms of grammar and sentence 

structure if it is used to polish an author’s own 

words. AI can expand the information available 

to authors in the literature and locate otherwise 

difficult-to-find sources, and it can be used to help 

initially develop a research idea. If AI is used in any 

part of the paper, the reviewer should also be aware 

and take the time to confirm the accuracy and any 

potential biases of any AI-based information in 

the article. AI should never be used in isolation 

to produce text without human oversight or input.

We discussed whether the use of AI should be 

acknowledged in a manuscript through in-text 

citations or in the acknowledgments section, or if 

it only needs to be reported through the journal 

submission portal. These discussions focused on 

the question of who benefits from knowing that 

AI was used, and why they need to know. For uses 

related to improving the author’s original writing, 

acknowledging AI software seemed unnecessary, 

and akin to acknowledging ubiquitous tools such 

as spell check within Microsoft Word. However, 

when the AI software was a critical component of 

the research, such as for image analysis, we deemed 

it necessary to acknowledge the AI software and 

version. Finally, because these are still early days 

for generative AI, we decided to include a question 

in the submission portal about AI use to better 

understand how often researchers incorporate AI 

in their manuscripts. This information would be 

used only for data collection and would not be 

passed on to the reviewers or editors.

We also considered the use of AI in code 

development. We determined that using AI tools 

to derive code is no different than adapting R 

code found online for a user’s specific purpose. 

However, while AI could be helpful in identifying 

holes or inconsistencies in a researcher’s code, it 

should not be used in stress-testing that code. We 

eventually agreed that AI-generated code can be 

used, provided that the authors acknowledge the 

AI assistance and detail its usage in the Methods 

section. An acknowledgment in the Methods 

section suffices if the AI was used for writing 

functions, adding documentation, or refactoring 

code for clarity. For example:

We used OpenAI's ChatGPT-4o to generate 

the initial implementation of the data 

processing function and to add inline 

documentation for improved readability.

These tasks are comparable to assistance 

gained through Google searches or consulting 

Stackoverflow, where authors remain responsible 

for the accuracy and correctness of the code. 

However, a detailed explanation of AI usage is 

required when AI is used to automate analyses, 

such as performing statistical analyses on 

tabular data (see https://help.openai.com/en/

articles/8437071-data-analysis-with-chatgpt). For 

instance:

We used OpenAI's ChatGPT-4o data 

analysis tool (gpt-4o-2024-05-13) to 

perform statistical analyses on our dataset, 

including generating summary statistics and 

visualizations. The AI tool›s methodology 

and output were reviewed and validated by 

the authors to ensure accuracy.

PSB 70 (3) 2024

213

In this example, the AI tool must be cited in a way 

that ensures the reproducibility of results because 

the AI significantly contributed to the analysis.

CONSIDERATIONS FOR 

REVIEWER GUIDELINES

Our committee also considered the use of AI in 

the review process. We decided that it does not 

help the journal or authors when the reviewer 

extensively uses AI to write their full review. The 

point of having peer reviewers is to obtain the 

researcher’s own unique expertise, which any 

General Author Guidelines

Use of artificial intelligence and large language models (generative AI):

Generative AI programs, such as ChatGPT, are widely accessible and commonly adopted across various 

scientific domains. When employing generative AI in scientific work, writing, or figure generation, it is 

crucial for authors to be aware that unintended content may arise, necessitating careful oversight. Authors 

must assume full responsibility for content produced by generative AI programs before incorporating it 

into the submitted manuscript.

Authors are requested to cite the use of generative AI when appropriate. For example, if generative AI is 

employed as an integral part of the methodology, it should be cited in the Methods section, specifying 

the manner of use, program, and version. The use of AI to address editing and proofreading does not 

require acknowledgement in the manuscript. Please see Wiley’s Best Practice Guidelines on Research 

Integrity and Publishing Ethics (https://authorservices.wiley.com/ethics-guidelines/index.html) for 

more information.                          

For Reviewers:

At (AJB/APPS/PSB), we highly value the professional expertise of peer reviewers to improve 

manuscripts published by the journal. Artificial intelligence (AI), including large language 

models or generative AI such as ChatGPT, is not allowed in the reviewing process. Uploading 

any author-submitted text, including the manuscript, abstract, or title, into an AI platform is 

considered a violation of confidentiality. The only exception is using AI as a tool to edit or 

proofread the language of a reviewer’s own work. 

Regarding Software and Code:

AI coding assistants have become increasingly powerful and commonplace. However, authors must be 

vigilant about the quality and accuracy of the generated code and take full responsibility for the results. 

Furthermore, authors who choose to use AI coding assistants are encouraged to take full advantage of 

their capabilities to generate tests, write documentation, and create robust, user-friendly, functional 

programs that can be more easily maintained and repurposed. In cases where AI is an integral part of the 

methods of the study, the authors should cite the program within the Methods section.

AI tool would lack. To abide by an AI program’s 

usage guidelines (such as for ChatGPT), reviewers 

should not input the manuscript or any part of it 

into a public AI tool because this would also be 

a breach of confidentiality. However, reviewers 

could potentially improve the spelling and 

grammar of their own written review using an AI 

tool, akin to a grammar or spell checker. 

Based on these conversations, our ad-hoc 

committee created AI policy for BSA journals as 

shown in the following box.

PSB 70 (3) 2024

214

ACKNOWLEDGMENTS

The authors thank Amy McPherson (managing 

editor of AJB), Beth Parada (managing editor of 

APPS), Richard Hund (production editor of AJB/

managing editor of PSB), and Emily Sessa (BSA 

Director at Large for Publications) for their insight 

during committee discussions and for their careful 

review of this paper.  

REFERENCES

Abramson, J., J. Adler, J. Dunger, R. Evans, T. Green, 

A. Pritzil, O. Ronneberger, et al. 2024. Accurate struc-

ture prediction of biomolecular interactions with Al-

phaFold 3. Nature 630: 493-500. 
Anderson, P. 2024. Academic writers on AI: An Oxford 

University Press Study. Website: https://publishingper-

spectives.com/2024/05/academic-writers-on-ai-an-ox-

ford-university-press-study/.
Authors Guild. 2024. Authors Guild demands prior 

consent for AI use of academic and news content. Web-

site:  https://authorsguild.org/news/ag-demands-prior-

consent-for-ai-use-of-academic-and-news-content/
Buriak, J. M., D. Akinwande, N. Artzi, C. J. Brinker, 

C. Burrows, W. C. W. Chan, C. Chen, et al. 2023. Best 

practices  for  using AI  when  writing  scientific  manu-

scripts: ACS Nanao 17: 4091-4093. 
Culley, T. 2018. How to avoid predatory journals when 

publishing your work. Plant Science Bulletin 64: 96-111.
Davis, P. 2023. Did ChatGPT just lie to me? The Schol-

arly  Kitchen Website: https://scholarlykitchen.sspnet.

org/2023/01/13/did-chatgpt-just-lie-to-me/
De Waard, A. 2023. Guest post – AI and scholarly 

publishing: A view from three experts. The  Schol-

arly  Kitchen Website: https://scholarlykitchen.sspnet.

org/2023/01/18/guest-post-ai-and-scholarly-publish-

ing-a-view-from-three-experts/
Dutton, C. 2024. Two major academic publisher signed 

deals with AI companies. Some professors are out-

raged.  The  Chronicle  of  Higher  Education Website: 

https://www.chronicle.com/article/two-major-academ-

ic-publishers-signed-deals-with-ai-companies-some-

professors-are-outraged?sra=true

Motoki, F., V. Pinho Heto, and V. Rodrigues. 2023. 

More human than human: Measuring ChatGPT politi-

cal bias. Public Choice 198: 3-23. 
Piers, C. 2024. Even ChatGPT says ChatGPT is racial-

ly biased. Scientific  American Website: https://www.

scientificamerican.com/article/even-chatgpt-says-

chatgpt-is-racially-biased/
Walters, W. H. 2023. The effectiveness of software de-

signed to detect AI-generated writing: A comparison 

of 16 AI text detectors. Open Information Science 7: 

20220158.
Wan, Y., G. Pu, J. Sun, A. Garimella, K.-W. Change, 

and N. Peng. 2023. “Kelly is a warm person, Joseph is 

a role model”: Gender biases in LLM-generated refer-

ence letters. Website: https://arxiv.org/abs/2310.09219
Weaver, W. N., and S. A. Smith. 2023. From leaves to 

labels: Building modular machine learning networks 

for rapid herbarium specimen analysis with LeafMa-

chine2. Applications in Plant Sciences 11: e11548.
Weaver, W. N., B. R. Ruhfel, K. J. Lough, and S. A. 

Smith. 2023. Herbarium specimen label transcription 

reimagined with large language models: Capabilities, 

productivity, and risks. American  Journal  of  Botany 

110: e16256.
Wood, H. 2024. Wiley and Oxford University Press 

confirm AI partnerships as Cambridge University Pres 

offers ‘opt-in’. The Bookseller Website: https://www.

thebookseller.com/news/wiley-cambridge-university-

press-and-oxford-university-press-confirm-ai-partner-

ships
Zhou, H. 2023. The intelligence revolution: What’s 

happening and what’s to come in generative AI. The 

Scholarly  Kitchen Website: https://scholarlykitchen.

sspnet.org/2023/07/20/the-intelligence-revolution-

whats-happening-and-whats-to-come-in-generative-ai/

PSB 70 (3) 2024

215

International Journal of 

Plant Sciences

IJPS 

is seeking contributions for a 

series of occasional papers, Primers 

in the Plant Sciences. “Primers” are 

short, peer-reviewed, accessible 

introductions to well-defined topics 

in the plant sciences.

Each Primer is both an introduction 

to a topic in plant science and a 

narrow-in-scope review that serves 

as a useful first-stop reference to 

scientists at all career stages. 

Primers are intended to provide the 

reader with a foundation in the topic 

and introduce them to leading 

research questions and 

methodologies in the field. 

Call for Proposals: Primers in the Plant Sciences

For more information, visit

journals.uchicago.edu/journals/ijps/primers

216

Bessey Award Winners Through the Years

As I started to near the end of my tenure as Editor-

in-Chief of Plant Science Bulletin, I thought a lot 

about what I wanted my last issue to include. 

One of my favorite parts of being editor has been 

exploring the archives and reading the words of 

contributors over the past 70 years. A prevalent 

theme that has run through the PSB  since the 

beginning has been botanical education. In fact, 

the very first article, printed on page one of the 

January 1955 issue, is an essay by the chair of the 

Education Committee, Sydney S. Greenfield. In 

his essay, Greenfield declares that Plant Science 

Bulletin will serve as a platform for discussions 

about education in plant science. 

“The Committee on Education of The 

Botanical Society of America has been 

studying means whereby it might effectively 

promote greater appreciation and proper 

development of plant science in the colleges, 

SPECIAL SECTION 

Honoring the Tradition of Botany 

Education in the 

Plant Science Bulletin: 

A COLLECTION OF ARTICLES BY CHARLES E. BESSEY 

TEACHING AWARD WINNERS

as well as the education of the general public 

as to the importance of plants and their study 

to man. It will require nationwide discussion 

among botanists of educational and other 

problems. with a view towards development 

and formulation of professional policies, and 

plans for coordinated constructive action.

Until now, a major obstacle to cooperative 

analysis and attempts to solve our common 

problems has been the lack of an appropriate 

medium for intra-professional discussions, 

and in this regard, the establishment of Plant 

Science Bulletin may well presage a new 

era for professional botany in this country.” 

(Greenfield, 1955)

The early editions of Plant Science Bulletin

are particularly rife with essays examining 

teaching philosophies and practices and setting 

out strategies for connecting with students, 

administrators, and the general public. I have 

found many of these articles, such as those by 

Palmquist (1956), Fuller (1957), and Stern (1971) 

to be especially impactful and I keep Palmquist’s 

Ten Commandments for the Teaching Botanist 

posted on my office door. The need for dialogue 

on educational themes, of course, endures and 

PSB contributors have continued to both debate 

educational ideals and share practical classroom 

activities (e.g., Wandersee and Schussler, 2001; 

Carter, 2004; Keller and Bordelon, 2022). I have 

used a variation of the Market Botany lab in my 

own botany course many times (Martine, 2011), 

By Mackenzie Taylor

Editor-in-Chief, Plant Science Bulletin

PSB 70 (3) 2024

217

and I am thrilled to have been able to feature 

many education-focused articles during my time 

as editor (e.g., Doust, 2016; Sundberg, 2016; 

Goodwillie and Jolls, 2018; Montgomery and 

Farrah, 2021; Parsley, 2021; and many others). In 

my last issue of Plant Science Bulletin, I wanted to 

showcase and further this long tradition.
It seemed obvious to me that this special feature 

could also provide a platform for some of our 

Charles E. Bessey Teaching Award winners. 

Fortuitously, many of these winners were already 

preparing talks for the symposium “Bessey’s 

Legacy: Enthusiasm and Innovation in Botanical 

Instruction,” moderated by Ben Montgomery and 

Rachel Jabaily at Botany 2024, and were willing 

to adapt these presentations into print essays. 

I reached out to other past winners, as well, and 

almost everyone graciously accepted my invitation. 
In my request, I asked only that contributors write 

about an issue of their choice having to do with 

teaching in botany. I suggested that articles could 

be a reflection on personal teaching philosophy, 

observations on the state of botany education, or 

a call to action for change. I’m pleased to say that 

the articles in this collection cover all of this and 

more. I found these articles to be inspiring and 

thought provoking and to provide a snapshot of 

the challenges and rewards of teaching botany 

in first quarter or so of the 21

st

 century. I firmly 

believe that Charles Bessey, as well as Sydney 

Greenfield, Edward Palmquist, and the other 

botany educators who have graced the pages of 

Plant Science Bulletin, would be thrilled to see that 

the members of the Botanical Society of America 

continue a strong a tradition of thoughtful and 

reflective teaching. 

REFERENCES

Carter, J. L. 2004. Developing a Curriculum for the Teach-

ing of Botany. Plant Science Bulletin 50(2): 42-47. https://

botany.org/userdata/IssueArchive/issues/originalfile/

PSB_2004_50_2.pdf  
Doust, A. 2016. RETurn to the Classroom: Linking Sci-

ence Teaching and Science Experience for Pre- and In-

service High School Science Teachers. Plant Science 

Bulletin  62(1): 25-29. https://botany.org/userdata/Is-

sueArchive/issues/originalfile/PSB_2016_62_1.pdf

Fuller, H. J. 1957. The Role of Botany in a Liberal 

Education.  Plant Science Bulletin 3(1): 4-6.  https://

botany.org/userdata/IssueArchive/issues/originalfile/

PSB_1957_3_1.pdf
Greenfield, S. S. 1955. The Challenge to Botanists. 1(1): 

1-4. https://botany.org/userdata/IssueArchive/issues/orig-

inalfile/PSB_1955_1_1.pdf
Goodwillie, C., and C. L. Jolls. 2018. Combating Plant 

Blindness and Plant Invasion Through Service-Learning. 

Plant Science Bulletin 64(1): 11-17. https://issuu.com/bo-

tanicalsocietyofamerica/docs/psb_64__1__2018
Keller, H. W., and A. Bordelon. 2022. Discovering the 

Microscopic World of Live Tree Bark: A Model Instruc-

tional Experience for Students and Teachers Using A 

Virtual iAdventure, Teacher Preparation Guide, Student 

Worksheets, and Moist Chamber Cultures. Plant Science 

Bulletin  68(1): 12-23. https://botany.org/userdata/Is-

sueArchive/issues/originalfile/WebPSB68_1_2022_3.pdf
Martine, C. T. 2011. Market Botany: A Plant Biodiversity 

Lab Module. Plant Science Bulletin 57(2): 61-66. https://

botany.org/userdata/IssueArchive/issues/originalfile/

PSB_2011_57_2.pdf
Montgomery, B. R., and K. P. Farrah. 2021. Teaching a 

Distance Botany Laboratory with Online, Outdoors, and 

Hands-On Exercises. Plant Science Bulletin 67(1): 16-28. 

https://botany.org/userdata/IssueArchive/issues/original-

file/PSB67_1_2021.pdf
Palmquist, E. M. 1956. Stimulation of Interest Among 

Undergraduates in Botany. Plant Science Bulletin 2(4): 

5-8. https://botany.org/userdata/IssueArchive/issues/orig-

inalfile/PSB_1956_2_4.pdf
Parsley, K. M. 2021. Plant Awareness Disparity: Looking 

to the Past to Inform the Future. Plant Science Bulletin 

67(2): 94-99. https://botany.org/userdata/IssueArchive/is-

sues/originalfile/PSB_2021_67_2.pdf
Stern, W. L. 1971. Responsibilities of Universities to 

Provide Trained Botanists for Undergraduate Education. 

Plant Science Bulletin 17(1): 2-3. https://botany.org/ps-

barchive/issue/1971-v17-no-1.html
Sundberg, M. D. 2016. Botanical Education in the Unit-

ed States. Part IV. The Role of the Botanical Society of 

America (BSA) into the Next Millenium. Plant Science 

Bulletin  62(3): 132-154. https://botany.org/userdata/Is-

sueArchive/issues/originalfile/PSB_2016_62_3.pdf
Wandersee, J. H., and E. E. Schussler. 2001. Toward a 

Theory of Plant Blindness. Plant Science Bulletin 47(1): 

2-9. https://www.botany.org/userdata/IssueArchive/is-

sues/originalfile/PSB_2001_47_1.pdf

218

Bessey Award Winners Through the Years

Four Things I Learned from 

30 Years of Teaching 

(That You Probably 

Already Know)

By Cynthia S. Jones

Department of Ecology and Evolutionary 

Biology, U 3043, University of Connecticut, 

Storrs CT 06250 USA

The Botanical Society of America is replete with 

excellent teachers.  Why? Because botanists have 

to be good teachers!  The inherent bias against 

plants in the United States virtually ensures that 

most students, at least until very recently, take 

their first plant class in college because it satisfies 

a requirement.  A good teacher erodes plant bias, 

ideally recruiting more than a few students to 

the “plant side.”  What we do as college teachers 

is incredibly important to ensuring the future of 

our academic discipline, and there is no better 

evidence than the BSA Membership Matters

survey (Figure 1) from June 2022. The majority 

responding discovered their passion for botany as 

undergraduates. I know I did.

When I first started teaching at UConn, I 

developed two upper division courses.  One 

was Plant Anatomy, which was already “on the 

books,” and the second was a course I called 

“Plant Developmental Morphology” based on the 

principles Don Kaplan taught at Berkeley (Kaplan, 

2022).  My approach to teaching was exactly what 

I had experienced as an undergrad and graduate 

student: a lecture/lab format where I gave the 

lecture, and a graduate student teaching assistant 

taught the lab.  Lectures involved chalkboards 

and switching between projecting Kodachrome 

images and an overhead projector. Lecture 

exams were structured in a short-answer format, 

primarily based on comparisons, descriptions, 

and definitions. Lab exams were based on moving 

from station to station every two minutes or so, 

largely focused on recognition of features.

I was not an early devotee of PowerPoint lectures, 

but three features eventually swayed me: I wouldn’t 

have to spend an hour before each lecture pulling 

Figure 1. Survey responses to the question “At what point 

did you choose botany as a focus of your career or inter-

est?” June 2022, BSA newsletter, Membership Matters.

PSB 70 (3) 2024

219

and organizing slides, I could incorporate new 

images much more easily, and finally, I would save 

myself the hassle of switching between overheads 

and Kodachrome slides.  Initially, my PowerPoint 

slides consisted of only images and graphs, but I 

soon discovered that using the chalkboard for the 

rest was awkward because when the projection 

screen was down, it covered the center of the 

chalk board.  It wasn’t long before I began to put 

the words and drawings I would have written on 

the board into the PowerPoint lectures.  Grabbing 

images directly from the internet also meant I 

could quickly incorporate new material without 

finding the original books and papers to photocopy 

for overheads. A win-win for me!

Within t

wo years, my evaluation scores declined, 

students seemed to sleep more in class, and they 

left the lab earlier and earlier. 

My realization that I was no longer providing 

students with a course that most of them valued led 

to some soul searching. For me it was a question 

of self-respect—if I was going be in the classroom 

(which was part of my job, after all), then I wanted 

to do what I could to make it go as well as possible. 

I’m not a funny person by nature, so I knew better 

than to try to motivate an audience with humor; 

nor am/was I brilliant enough to captivate students 

just by talking without much prep.  My approach 

was to present a course focused on content that 

was as clear as I could make it and by building a 

story line that flowed so one bit of information led 

to another.  For me, plant development was the 

obvious thread, so both of my courses began with 

embryos and the plant body grew from there. 

All this soul searching resulted in two insights: 

(1) I’m introverted enough that although I loved 

teaching, I never really “enjoyed” giving lectures—I 

learned to do it well eventually—but what I loved 

were the students’ “ah ha” moments in the lab, and 

(2) I was drawn to teaching because of the plants. 

I had to figure out how to let the plants lead.  I 

decided to try to restructure my courses around 

Don Kaplan’s mantra “Ask the Organism

.”

1. Ask the organism

In 2017, my then PhD student Dr. Kerri Mocko, 

who had been my TA for several semesters, 

graduated.  Before she left for her post-doc, I was 

able to pay her as an adjunct for one semester and 

together, we overhauled Plant Anatomy into what 

we called Plant Structural Diversity. 

1. First, we created a “studio” time slot. We 

changed the schedule from one three-

hour lab to two two-hour labs per week, 

with the lab scheduled to directly follow 

the “lecture,” resulting in two three-hour 

time blocks.

2. We emphasized at the beginning of class 

that we were teaching a “skills” course, 

not a course that required memoriza-

tion, but at the same time stressing the 

importance of the vocabulary.  We told 

students the first day that by the end 

of the course, we expected them to be 

researchers, in that they should be able 

to make a hand-section of any vegeta-

tive organ on most plants and explain to 

someone else its internal structure and 

function of cells.  Exams would be based 

on interpreting material they had never 

seen before, or interpreting something 

they had seen during the course, but 

from a different perspective.

3. We revised every lecture (“content de-

livery”) so that it emphasized structure, 

function, and evolution together (e.g., 

a simplified phylogenetic approach to 

wood structure was followed immedi-

ately by ecological wood anatomy).

4. We reorganized the lab manual so that 

rather than being written in paragraph 

form (because students didn’t seem to 

read carefully enough to figure out what 

they should see), the text was largely 

structured in bullet points with open 

boxes where they should draw specific 

features. Consequently, the important 

parts of what they should take away from 

each unit were abundantly clear.

5. We carefully matched the lecture and lab 

material in short time units, so that one 

of us would present content (lecture) for 

10–25 minutes (at most, with a few ex-

ceptions), and then students would turn 

to the “active engagement” exercises (i.e., 

PSB 70 (3) 2024

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the lab material).  When they finished 

looking at the material (we always asked 

if it was okay to move on, or we had 

them put bright sticky notes on their mi-

croscopes to indicate they were finished), 

we moved into the next content delivery 

section.  This approach had numerous 

advantages: 

a. Switching between listening and 

active engagement keeps students 

awake!

b. We could stop talking anytime so 

students could make hand sections 

and see for themselves what we had 

just been talking about. This immedi-

ate reinforcement with living material 

turned out to be a powerful teaching 

tool.

c. In previous years, we put all the plant 

material at the back of the room 

and had students pick up what they 

needed once lab began. Now, we put 

as much of the live material on their 

tables at the beginning of class as pos-

sible, so when they sat down, it was 

right in front of them.  This had two 

advantages: (1) some students started 

looking at the material when they sat 

down, rather than their phones and 

(2) we could begin the unit with a 

question about the material in front 

of them; in other words, as much as 

possible we asked them to “Ask the 

Organism.”  

d. Students didn’t finish units at the 

same rate. We encouraged anyone 

who was done to move around, go 

outside for a few minutes, etc. What 

happened most often was that stu-

dents would help each other or just 

start chatting. At first, I was a little 

dismayed that they were talking about 

anything but what they were looking 

at, but I soon realized that if we didn’t 

interfere, the conversation slid easily 

between a show they’d just watched to 

“Is this the secondary cell wall?” and 

then back to the show.  We tried to 

keep the chatter at low volume and as 

far as I know, no other students com-

plained that they couldn’t concentrate.  

Encouraging the movement, breaks, 

and the social aspect helped boost the 

general enthusiasm and energy level 

during the long afternoons.  

6. Lecture and lab exams were not separate.  

Instead, on the day of the exam, students 

would arrive to find a few plants (or 

slides) at their desk that they may or may 

not have seen before. If a living plant was 

represented, they would be responsible 

for making hand sections. The exam 

would consist of four or five questions 

that would require illustration, interpre-

tation, and the rationale for their inter-

pretation; most students stayed the full 

three hours to complete the midterms. 

From semester to semester, we tried 

letting students bring in references, i.e. 

their notes.  They were not allowed to use 

the internet, though.  As far as I can tell, 

allowing notes didn’t really affect their 

grades much, but it did seem to reduce 

the stress associated with taking exams.  

Another thing I’ve done since before we 

restructured the course is that I would grade the 

first exam, and then give everyone in the class 

the chance to redo it as a take-home, re-grade 

the take-home exam, and then record the average 

of their scores.  I emphasized to them that my 

concern was for their learning, not their grades.  

In over 30 years of teaching, I had only one student 

complain to me that this approach wasn’t “fair” to 

the good students.  I pointed out that I don’t grade 

on a curve, so the “mean” (and the comparison 

it implies) was not relevant. (I wanted students 

to know how they are doing as we went through 

the course, and not count on a mystical curve 

at the end to save them.)  Students who scored 

well the first time didn’t need to spend the extra 

time redoing the exam.  I also pointed out that I 

generally only do this for the first exam.  From 

my perspective, it helped the students who didn’t 

do well on the first exam maintain some degree 

of interest in the course, without feeling like there 

was no “hope” of eventually getting a good grade.  

This perspective is completely selfish on my part, 

because who wants to spend the semester trying to 

teach students who don’t want to be there because 

they have no hope of attaining their goals?

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221

Student reaction to the integrated lecture/lab 

format was strongly positive, so the following year, 

I reorganized Developmental Plant Morphology 

to follow the same format.  Since that time, a 

few other colleagues in my department have 

restructured their lecture/lab classes in a similar 

format as well.

Does it take more time?  Probably.  I arranged the 

rest of my schedule so that as much as possible, I 

devoted two full days a week to teaching, but very 

little time the rest of the week. And since I was in 

the classroom during the lab, I no longer needed to 

spend a few hours each week on TA meetings.  The 

TA and I set up the lab together and I explained 

what I wanted students to see during that process.  

In order for the TA to gain more experience 

being in control of the classroom, I offered the 

TA the opportunity to be the lead instructor, i.e., 

providing the content and leading the engagement 

periods, on as many units as they wished.  Most 

TAs were happy to lead one or two days out of the 

semester but didn’t really want to take the lead 

more than that.  Was the extra time worth it?  For 

me personally, absolutely! This format seemed 

to make it easier to spark a deeper interest in 

the material. I felt like I got to know my students 

better, which helped me better understand their 

questions and responses to the material, and I 

ended up with additional insight into what aspects 

of the course worked well.
2. Students like drawing
The traditional approach to recording information 

in visually oriented courses (comparative anatomy, 

morphology, etc.) is drawing, but at some point, 

I began to worry that students would prefer to 

work in a digital format, i.e., that my approach 

was old fashioned and didn’t involve the “latest 

technology.” More and more students were using 

their phone to take pictures through the ocular of 

the microscope.  While I’ve seen some amazing 

photos taken this way, getting the focus right 

requires very steady hands and careful positioning. 

To try to make this easier, in 2015, I received a 

teaching grant from my university to purchase a 

large screen (since technology in the old building 

I taught in was rudimentary) and ten iPad minis 

(i.e., one for each pair of students).  I worked 

with the UConn’s Technical Services department 

to design adapters that would position the iPad 

minis in place over one ocular for accurate focus.  

Now, for the first time, individual students could 

share though Airplay what they were seeing on 

their scope to the large monitor so that everyone 

could see it.  

I also posted PDFs of each lab write-up so students 

could populate the lab write-up with digital photos 

that could be labeled in OneNote.

Surprisingly, almost all students preferred to draw 

on the lab handouts.  For one thing, the iPad 

adapter ended up being slightly cumbersome 

to set up and remove, and since it blocked one 

ocular, students preferred not to keep it mounted 

on the scope. More importantly, students also told 

me that especially since the pandemic and flipped 

classrooms, almost all of their education was 

digital.  They spent hours upon hours each day in 

front of screens, so it was actually a relief to come 

look at something and try to draw it.  

3. Students want to do well, but life gets in the way

The first decade of my career I was always on the 

watch for people cheating, wary of those trying 

to take advantage of the system, and of me. I 

was skeptical of excuses. I became much more 

sympathetic, and I think a better teacher, when 

my step-kids became university students. I finally 

began to understand the student experience 

from the student perspective.  If students missed 

assignments or didn’t perform well on a test, I 

could appreciate that they had other things going 

on in their lives besides my course.  Almost 

all were pursuing minors, double majors, and 

multiple club or organizational activities, in 

large part responding to stress of building their 

resumes. After I switched to the integrated lecture/

lab structure, I also learned just how many of my 

students went from my class directly to a job.

PSB 70 (3) 2024

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Many people have suggested that course 

evaluations from students don’t reflect student 

learning.  Perhaps… but evaluations often reflect 

how students feel about a course.

Students might not remember the details of what we 

tried to teach, but they will remember how they felt 

about a class and by extension, the subject matter.  

More than anything else, my goal as an educator 

was to leave students with an appreciation (dare I 

say love?) for plants and how they grow, to teach 

through awe and discovery as much as possible, 

and to respect them as people.

Are there things I wish I’d done differently? 

Absolutely.  In hindsight, I wish I had incorporated 

more inquiry-based techniques than I did. Even 

so, nearly all students reported on evaluations 

that they learned more, or much more, than 

in their other classes. One thing that worked 

well was to conduct scavenger hunts inside our 

UConn Botanical Conservatory (aka, The EEB 

Greenhouses).  Scavenger hunts proved to be 

effective teaching tools because students had to 

use their skills to rule out possible candidates, 

as much as to investigate those that exhibited a 

feature of interest.  I recognize that an incredible 

collection of plants in greenhouses adjacent to 

our teaching lab building has been an incredible 

privilege. It also has been one of the greatest joys 

of my career. 

4. Spend more time outside

Despite our amazing indoor plant collection, 

I regret that I didn’t have students spend more 

time outside in both plant structural diversity 

and developmental plant morphology. Over the 

last two decades, many studies have shown that 

outdoor classrooms and educational activities are 

incredibly effective for increasing concentration, 

creativity, and retention in children (e.g., Coyle, 

2010; Kuo et al., 2017).  The benefits of outdoor 

classrooms at the college level are understudied 

(Birdwell, 2024), but based on feedback from 

students, the few units of my courses that 

did require spending time outside (e.g., tree 

architecture) were always the most popular. 

Some of the scavenger hunts in the greenhouse 

could be converted to outdoor activities, weather 

permitting, if I had made the effort.  I wish I had.
I have come to believe it is possible to teach our 

traditional courses like plant anatomy in such a 

way that students not only come to appreciate 

plants, but also develop an understanding, even 

if subconscious, of a plant’s place in the world, 

while at the same time deepening their own con-

nections to the natural world. Would it take some 

effort to revise my courses again to do so? Abso-

lutely. But at a time when over 80% of Americans 

live in urban environments, it feels imperative 

to give it a try.  Hopefully, future generations of 

botanical educators have already, or will tumble 

to this realization sooner than I did.

REFERENCES

Birdwell, T., M. Basdogan, and T. Harris. 2024.  Devel-

oping outdoor campus space for teaching and learning: 

a scoping review of the literature. Learning Environ-

ments Research DOI: 10.1007/s10984-024-09504-1. 
Coyle, K. J. 2010. Back to school: back outside! How 

outdoor education and outdoor school time create 

high performance students. A report from the National 

Wildlife Federation. Website: https://www.nwf.org/Ed-

ucational-Resources/Reports/2010/09-01-2010-Back-

to-School-Back-Outside
Kaplan, D. R. 2022. Kaplan’s Principles of Plant Mor-

phology. CRC Press, Taylor and Francis Group, Boca 

Raton, FL.
Kuo, M., M. Browning, and M. L. Penner. 2017: Do 

lessons in nature boost subsequent classroom engage-

ment? Refueling students in flight. Frontiers in Psychol-

ogy 8: 2253. 

223

Bessey Award Winners Through the Years

By J. Phil Gibson

Professor, School of Biological Sciences

University of Oklahoma

My colleague Dr. Drew Hasley recently began a 

seminar presentation with what appeared to be 

a blank, white screen, and he asked the audience 

to read what it said. Although we were unable 

to read the text on the slide, the software on 

Drew’s computer—something he uses daily to do 

all the things we think of as the “typical work” 

of a scientist—was able to detect and read the 

title of his talk. The title was written in a white 

font on a white background, and Drew, who is 

blind, explained that font color and background 

are irrelevant to screen readers. He then said, 

“People are not disabled. It’s environments that 

are disabling.” Following this incredibly powerful 

combination of demonstration and statement, he 

proceeded to describe what can and must be done 

to improve accessibility to STEM education in our 

classrooms, lecture halls, and laboratories. 
The question, the challenge, the hurdle that 

undoubtedly arises for many of us when we 

consider accessibility for our science classes 

is: “How can I adjust my teaching spaces and 

practices to be more inviting and provide 

opportunities for all students to learn?” There is 

no one-size-fits-all answer to that question. Some 

solutions may be relatively simple, such as giving 

extended time on assignments. Others may be 

more challenging to discern and implement, such 

as making a laboratory activity compliant with the 

Americans with Disabilities Act. Regardless of the 

complexity of the problem, Universal Design for 

Learning (UDL) can be an effective starting point 

to help teachers identify strategies and solutions 

to support learning for all students and help them 

achieve their goals as a scientist regardless of a 

student’s visible or hidden disabilities. 

Universal Design for 

Learning Botany

UDL is based on the architectural principle 

of Universal Design, in which constructed 

environments intentionally contain design 

features to improve accessibility or use for one 

group of people that absolutely needs that specific 

design element and can also provide emergent 

benefits to others. The classic example is a feature 

we regularly see on sidewalks called “curb cuts.” 

These small ramps are commonly found in curbs 

and at intersections. They are essential, and 

required by law, to help individuals with mobility 

issues use wheelchairs or walkers to safely 

navigate sidewalks. However, they also benefit 

people pulling luggage, pushing a cart, or many 

other activities where the curb is anything ranging 

from a nuisance to a literal barrier. Automatic 

doors provide a similar benefit. Some people need 

them, while others benefit from their availability. 

UDL is based on the same idea. By designing 

and providing educational experiences that 

intentionally include features essential for some 

students to use them, we can also provide both 

anticipated and unanticipated learning benefits to 

everyone in the classroom. 

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UDL originated with the educational non-profit 

group CAST. Their mission for over 40 years has 

been to promote learning spaces and experiences 

that “are intentionally designed to elevate 

strengths and eliminate barriers so everyone has 

the opportunity to grow and thrive” (CAST.org). 

Their approach prompts educators to recognize 

that there is no “typical” student and that the real 

“normal” situation in any classroom is a range of 

differences among our students in how they learn 

and express what they have learned. UDL breaks 

these differences into three categories: Engagement, 

Representation, and Action/Expression. CAST 

researchers have shown that these areas are 

fundamental to how learners interact with lessons, 

perceive and take in information, and demonstrate 

their understanding, respectively. For each of these 

categories, there are three elements: Accessing 

(how students can obtain and use information 

and resources), Building (how students construct 

knowledge, skills, and understanding), and 

Internalizing (how students reflect upon, apply, 

and retain learning).  CAST combines the three 

categories and their three elements into the UDL 

guideline matrix (https://udlguidelines.cast.

org) to give educators suggestions for modifying 

lessons and removing unnecessary barriers 

to make learning opportunities available and 

meaningful to all students.
For example, suppose you invite a speaker to 

your class, but you have a student who is hearing-

impaired. The challenge in this situation lies at the 

intersection of Representation and Access in the 

UDL guidelines matrix. A suggested solution is to 

provide an alternative means of representing what 

the speaker is saying, such as providing a real-

time transcript or an American Sign Language 

(ASL) interpreter. While these items are essential 

for our hypothetical student, the transcript could 

also benefit students seated in the back or in 

a noisy part of the room, or perhaps someone 

who missed the lecture. The interpreter could 

potentially even benefit students studying or 

who know ASL. Likewise, the experience could 

increase awareness and stimulate interest to learn 

ASL.  Providing collections of slides before class 

is another example. Some students may have an 

accommodation that requires providing lecture 

slides to them beforehand, but providing them to 

all students can benefit others as well. The reasons 

of how or why they could benefit other students 

is irrelevant. Unless there is a specific reason that 

sharing them would somehow hinder learning, 

why not give everyone the benefit of having the 

resource?
Other challenges require more complex solutions. 

For example, my classes frequently involve 

constructing and interpreting phylogenies. For 

a blind or low-vision student, learning these 

things is an immense challenge. Through my 

collaboration with Drew Hasley, Kristin Jenkins, 

and Hayley Orndorf, we modified an existing tree-

thinking resource called the Great Clade Race (1) 

that uses symbols printed on cards and is therefore 

dependent on vision to teach tree thinking by 

converting it into one that uses tokens, making it 

a Tactile Clade Race (2). The tokens are accessible 

both visually and tactilely, so just like a previous 

example, different forms of representing the 

information increased accessibility by removing 

the barrier of vision-only access. All other 

elements of the activity remained the same, and 

assessments demonstrated its effectiveness at 

teaching the concepts and skills (2). However, we 

also noticed that in the tactile version, students 

worked as teams and completed the activity faster 

than for the visual form. This revealed several 

unintended and unanticipated benefits of our 

modification. And therein lies the true value of 

UDL: it stimulates changes that are essential for 

some but beneficial to all. 
UDL is neither a step-by-step process nor a 

curriculum structure one can follow to make 

lessons and resources accessible. Rather, UDL is 

a set of guidelines and suggestions for considering 

and identifying items to change. It should be 

thought of as an approach or perspective rather 

than a checklist to solve problems. Through 

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225

thoughtful consideration of course goals and 

learning objectives, UDL can help frame issues 

and identify solutions. For example, essay 

questions are common components of exams and 

other assessments to evaluate learning. However, 

consider whether an essay is the only way a student 

can express understanding. When grading an 

essay, ask yourself if you are also using their writing 

and grammar skills as indicators of understanding 

the topic in the question. Those are two different 

skill sets. If writing is part of the assignment, 

learning goals, and rubric, there is nothing wrong 

with evaluating writing itself—but what if your 

goal is to determine if the student understands 

a botanical concept such as the structure of a 

flower? Would a diagram be an acceptable way 

to demonstrate learning as well? What if they 

are not a particularly strong wordsmith, but they 

are excellent at producing diagrams? We have all 

told students at one time or another that we are 

not grading their artistic skills when we ask them 

to produce labeled diagrams in assessments. 

But how often do we extend that same leeway 

to questions when, for no specific pedagogical 

reason, we ask for or expect written answers by 

default or because that’s an easier question to 

write? Providing multiple, appropriate means of 

expression to show understanding is a solution 

at the core of UDL. If you are hesitant about that 

suggestion, consider this. In a recent workshop, a 

UDL expert demonstrated how a UDL perspective 

can help us better promote and evaluate learning. 

Here is a botanical modification of their activity to 

demonstrate this point. Get a pen and paper and 

draw a flower. First use your dominant hand. Do 

the same using your non-dominant hand. Now 

hold the pen or pencil in the crook of your arm 

or with your foot to draw a flower. Now suppose 

I am evaluating your knowledge of botany 

based on whether you drew the parts correctly 

and how well you drew a flower with your foot. 

Although some of you may draw quite well with 

your foot, that would hardly be a fair assessment 

of knowledge, right? I may have a perfectly valid 

reason for trying to find someone who can best 

draw a flower with their foot. But if what I want 

to determine is whether you know the parts of a 

flower and how they are put together, why would 

I try to base my evaluation of knowledge on the 

quality of the drawing? We make a similar mistake 

when we expect students to demonstrate thoughts 

and knowledge in a restricted way that may not 

allow them to be at their best to express them. 

When we use writing skills to evaluate knowledge 

of something else, we are making the same error 

as in my flower-drawing example. UDL provides 

ways to prevent that from happening. I am not 

advocating that we let students decide which 

assignments they will do or the form they will take 

for all assignments, although that is an intriguing 

idea. What I am asking you to consider is whether 

there are other ways, or more available  options, 

that would allow students to do their best work. 
There might be one question remaining that you 

are asking about UDL: “Why do this before I 

need to do it?” Of course, we would all provide 

any accommodations for students upon being 

informed by the appropriate campus office, and 

so one could wait until the need arises to do any 

of this. To that point, I ask that you consider 

the differences between accommodation and 

accessibility, and the consequences of their 

differences on teaching and student experiences. 

Both share the important goal of increasing 

inclusion of all learners in the classroom or 

laboratory. However, accommodation typically 

involves reacting to specific needs once made 

aware of them—usually just before the start 

of a semester only a few days away—and then 

modifying lessons, activities, or assessments so 

they can be used by a particular learner. This often 

results in frantic, last-minute changes at a time 

when there is little free time available. In contrast, 

thinking about how one can improve accessibility 

via UDL takes a proactive approach by carefully 

considering, designing, or modifying lessons 

and activities from their inception or as part of 

regular curricular updates to include features that 

are essential to support learners that have specific 

needs and can potentially benefit everyone. 

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This approach to solving problems beforehand 

usually results in more thoughtful solutions 

that are aligned with learning goals rather than 

last-minute modifications that just need to be 

“good enough” to work. A proactive accessibility 

stance is better than a reactive accommodation 

stance for several reasons. It increases inclusivity 

by ensuring that learning environments and 

experiences allow everyone, regardless of ability, 

to participate fully. It is more efficient simply 

because including accessibility features upfront 

saves the instructor time as compared to adjusting 

later. Anticipating learning challenges and student 

needs is also empowering for individual students 

because it gives them the freedom to navigate 

their learning and opportunities to do so without 

asking for special accommodations. Last, and 

possibly most importantly, using a UDL stance 

to increase accessibility benefits for everyone in 

the classroom or lab creates a positive learning 

and working environment because it promotes a 

culture of inclusion and respect for all students 

and their needs. 
Albert Einstein once said, “Everyone is a genius. 

But if we judge a fish by its ability to climb a tree, 

it will live its whole life believing that it is stupid.” 

We should think about that before we step into our 

classrooms. We often teach and assess in ways that 

are comfortable to us, ways we have experienced, 

ways we would show our understanding, or ways 

that we haven’t really dissected pedagogically. 

It’s easy to think that if it worked for us, it must 

be good—or at least it will work for the average 

student. As I mentioned earlier, the average 

student is a mythical creature. Our students are a 

rich tapestry of diverse needs, experiences, goals, 

abilities, and motivations. We must remember that 

what we do in the classroom or laboratory is about 

providing students experiences and opportunities 

to learn, gain skills, develop skills, and do their 

best. I am not asking anyone to immediately make 

wholesale changes in their teaching. I am asking 

that we all at least examine our classes through a 

UDL lens, and identify one thing or one aspect of 

a course or a lesson that can be improved by UDL 

modifications. Doing that one little thing can have 

a huge, positive impact. As botanists, we are quite 

familiar with that idea. Just remember that doing 

one little thing to increase accessibility is just like 

planting a seed. And we all know how the one 

small action of planting a seed, like knowledge, 

can have huge consequences once it starts to grow. 

REFERENCES

Goldsmith, D. W. 2003. The Great Clade Race: Pre-

senting Cladistic Thinking to Biology Majors & Gen-

eral Science Students. The American Biology Teacher 

65: 679–682.
Hasley, A. O., K. P. Jenkins, H. Orndorf, and J. P. Gib-

son. 2024. Tactile Trees: Demystifying Phylogenies 

for Everyone with Universal Design for Learning. The 

American Biology Teacher 86: 281–288.

227

Bessey Award Winners Through the Years

By Christopher T. Martine

Department of Biology, Bucknell University

A few days ago, I had a visit with my Field Botany 

class to the campus farm on the southeastern 

edge of Bucknell University, where I have been 

employed as a professor since 2012. The visit to 

the farm, where we met crop plants and discussed 

their taxonomic connections to the wild species 

we have thus far learned, capped off a big week for 

our group. For the previous class we left campus at 

7:30 a.m. for a 4-hour trip to the Mohn Mill Natural 

Area, a designated Wild Plant Sanctuary in Bald 

Eagle State Forest. The site, just off Pennsylvania’s 

Mid State Trail, is dotted with large circular vernal 

pools populated with enough “Osmunda” ferns 

to make one feel as though you have stepped 

back into the Jurassic. But the highlight of this 

annual excursion is always the moment when 

I tell the students, as they are lined up across a 

narrow wooden bridge on the Mid State looking 

down into the forest, that—only 5 weeks into the 

course—they would be hard-pressed to find a tree, 

shrub, forb, or fern that they don’t recognize and 

know the Latin name for (Figure 1). 
This group of 17 juniors and seniors had learned 

something like 75 species by that point, so their 

handle on this particular woodland was a function 

of similar plant communities we’ve visited and 

the plants we’ve seen in them (and, of course, the 

work the students have put in to memorize Latin 

names and recognize species when they come 

into view). Still, even with caveats, staring into 

nature and realizing you are seeing it in an entirely 

different way than a month ago is a powerful and 

deservedly pride-inducing moment. 

Field-Based Courses Still 

Matter, but not Like 

They Used To

So who are these students? Almost all of them, 

like many of the students I have taught in 19 years 

of professing at Bucknell and SUNY Plattsburgh, 

are Biology majors taking my class to fulfill an 

upper-level requirement in ecology/evolution. 

Many of them are headed for careers in health and 

medicine; a number of them will be MDs. Content-

wise, Field Botany is a one-off for the majority of 

my enrollees. And, if I am being honest, this is 

one of the best things about teaching the class: 

it is a one-semester opportunity to initiate a life-

changing shift in perspective. 
When I took my first field course, Dendrology, at 

Rutgers University in the fall of 1993, this is exactly 

the shift that happened for me. I added a few 

more undergrad course-based field experiences, 

thereafter, including two 5-week summer field 

courses in Newfoundland and in Alaska. But, for 

me, the die had been cast the moment I aced my 

first tree/shrub identification quiz. I knew that I 

would someday seek out opportunities to teach in 

similar ways, with the hope and intent to connect 

students to nature by teaching about real things in 

real places. 
I started as an undergrad TA in that same Dendro 

course; later, as a Master’s student, I taught the 

whole class as a sabbatical replacement. Working 

PSB 70 (3) 2024

228

for two government agencies I managed K-12 

outdoor education programs and then, privately, 

co-ran a few years of K-4 summer outdoor camps 

with my wife, Rachel. And when I got my first 

tenure-track job at SUNY Plattsburgh in 2006, I 

immediately added Field Botany to the Biology 

curriculum.  At that point in time, the students I 

was teaching had had childhoods a lot like mine. 

Students who were 21 or 22 years old in 2006 were 

born in the mid-1980s; they had dealt with idle 

time free of constant smart phone access. They 

were inherently aware of their surroundings much 

of the time. 
Yet by the time I arrived at Bucknell in 2012 and 

developed a new version of Field Botany, I could 

already tell things were changing. So could a 

lot of people, as perhaps best evidenced by the 

publication and popularity of Richard Louv’s Last 

Child in the Woods in 2013—and the suddenly 

widespread use of the phrase “nature deficit 

disorder.” 
Some would say things have only gone downhill 

since then. Students who are 21 or 22 years old in 

2024 were born around 2003. Everything about 

their childhoods was different from mine. Smart 

phones, tablets, laptops, constant connection… 

and constant reasons for not being outside, or 

taking long hikes, or camping or fishing or looking 

up at the night sky. These students grew up over-

scheduled, over-managed, and overly focused on 

extracurriculars and youth sports. Many spent 

years of their lives outside—but on athletic fields, 

not in the woods or down in the creek. Their 

connection to nature has been mediated by screens 

or experienced through enough filters to make 

nature itself feel like artifice. The distance between 

students and a comfortably broad understanding 

of the biodiversity around them persists even after 

heading off to university. 
Upon arrival to college, Biology and similar 

majors are often now plugged into courses with 

integrative approaches that have become the way 

we introduce biology at many institutions, with 

current students not experiencing (for better or 

worse) the same semi-exhaustive march through 

general biology content that previous generations 

faced. Case in point: Bucknell’s new four-course 

introductory core sequence. Lauded on campus 

for an approach focused on student retention, 

skills-building, and accessibility, our content-

based courses consist of case studies that vertically 

integrate subdisciplinary content. What was once 

a 4-week unit on plant diversity and evolution 

is now a 4-week module on “Milkweeds and 

Monarchs” touching on topics including ecological 

niches, interspecific competition, plant response 

to herbivory, transport across membranes, neuron 

structure and function, impact of mutations, 

and predator adaptation. It’s all pretty great and 

students gain a lot with this approach, learning to 

understand the multi-dimensionality of biological 

problems. But one trade-off is that they also 

receive less traditional content, including a deep 

primer on general botanical concepts. When these 

students arrive as juniors or seniors in my Field 

Botany class, they typically know little about the 

life of plants unless they have learned it outside of 

their formal education.
“Get off my lawn/front-yard-wildflower-meadow,” 

cries the old-guy botanist. 

Figure 1. Students enrolled in the 2024 edition of Field 

Botany at Bucknell University on Pennsylvania’s Mid 

State Trail, one of 20 locations the class will visit this se-

mester. 

PSB 70 (3) 2024

229

I am (in this moment) not being that guy, however. 

Because as much as the above items do present 

a challenge, they also present an important 

opportunity. 
For proof, let’s return to the Mohn Mill Natural 

Area, where my class spent time hanging 

around a sphagnum-dominated mountain 

“boglet” discussing glacial cycles and rates of 

decomposition. A student later reflected that 

they had heard about bogs in other classes, and 

even learned the story of “Tollund Man” (the 

ancient preserved body recovered from a bog in 

Denmark), but they never imagined that they 

might live anywhere near a similar sort of place—

let alone one they would someday visit IRL. To 

be standing in an actual “bog,” feeling the give of 

the peat, allowing the water to infiltrate your old 

pair of sneakers and soak into your socks… to be 

introduced to plants that grow nowhere else but in 

these particular habitats and to understand why… 

this completes the picture. This is the stuff you 

now never forget. 
When we teach field courses, we provide the 

context to so much of what our students have 

already learned and may learn later on; it is 

integrative biology on steroids. 
Increasingly, these courses are also providing the 

first real opportunities for students to experience 

nature in meaningful ways. As a baseline, even 

before the content delivery and the graded 

assessments, this is already enough to change the 

way a person feels in the world and to spark an 

appreciation for the life around them. This is why 

field-based courses still matter, but not like they 

used to. These days, they might just matter more 

than they ever have. 

230

Bessey Award Winners Through the Years

A key feature in connecting students to nature 

and the botanical world is to get them to “see” the 

diversity of botanical textures in the green world 

around them. Neo-natural history, where we take 

a closer look at each plant, adds new dimensions 

and wonder at the way we look at plants. It trains 

the eye to recognize the importance of plant 

diversity in scientific discovery and to understand 

the critical role of plant diversity on our quality of 

life. If we are to make progress in addressing the 

two main environmental crises of climate change 

and biodiversity loss (Figure 1; Pörtner et al., 

2023), an appreciation and understanding of the 

multiple ways plants provide solutions is critical. 

Plants are key in solving these dual environmental 

crises (Griscom et al., 2017; Pörtner et al., 2023; 

Zielinski et al., 2023).  There is no more important 

time than now for us to teach plant diversity both 

to make new discoveries and to find creative 

solutions to environmental problems.
Seeing the diversity of plants first hand in the 

field and lab provides an indelible experience that 

trains the eye to see, leads to new discoveries, 

provides examples of the power of the comparative 

approach, and gets students to recognize the 

critical role of biodiversity in our environment.  

With over 400,000 plant species globally (Enquist 

et al., 2019), we as botanists are blessed with 

having an almost limitless number of species to 

explore and to fuel new discoveries.  Almost every 

plant has something unique or special.  From the 

seemingly simple morphology of bryophytes to 

the extraordinary diversity of the most complex 

flowers, fascination dominates.  Here I highlight 

Neo-Natural History:  

Careful Observation and 

Co-Discovery in Teaching 

Botany

By Joan Edwards

Williams College

Figure 1.  Plant diversity can provide solutions to the 

twin environmental crises of climate change and biodi-

versity loss, which are coupled through human-caused 

dynamic interactions.  Each of the three factors on the 

triangle (biodiversity, climate change, and society) im-

pact each other (red arrows).  Humans have the ability 

to improve our quality of life by mitigating the negative 

impacts (blue arrows) and in return, restoring, or gaining 

valuable services (green arrows). (Adapted from Pörtner 

et al., 2023.)

PSB 70 (3) 2024

231

two approaches. The first is to look in depth at 

multiple features of a plant to highlight how they 

persist and their role in the environment.  The 

second is to look at a plant over time to chart the 

evolution of plant behavior.  I describe easy-to-

access examples that may surprise, delight, and 

inform.  
Looking closely at the multiple dimensions of 

a single plant creates a fuller picture of plant 

function and its role in the environment.  An 

excellent example is close examination of 

Marchantia (Figure 2A), a relatively easy to access 

liverwort often growing at the base of buildings 

(even in February), but also found as a “volunteer” 

in greenhouses.  
These small non-vascular plants are worth a close 

look because they are relatives of the first land plants 

(Qiu et al., 1999) and had a profound impact on 

the environment.  These early cryptospores were 

responsible for lowering the atmospheric levels 

of CO

2

 and for triggering a mini–ice age during 

the Ordovician over 400 mya (Lenton et al., 2012).  

They are also responsible for the current levels of 

oxygen in our atmosphere (Lenton et al., 2016). 

There is almost a disconnect when students realize 

these small, seemingly inconspicuous plants had 

such a profound impact on our environment.  If 

small liverworts can impact climate, what about 

the impact of eight billion people?   
If we look at Marchantia’s relatively simple 

morphology (Figure 1A), we see a dichotomously 

branching ground creeper that can never grow 

tall but can hopscotch across the landscape by 

harnessing the power of raindrops to jettison 

gemmae to new locations. The gemmae cups 

(Figure 2B) provide an example of evolutionary 

design where the urn shape provides a lower 

chamber in which new gemmae are produced and 

the top funnel shape serves as a launching site to 

Figure 2. (A) Top view of the dichotomously branching liverwort,

Marchantia,

 with mature and developing gemmae 

cups extending from the surface.  Individual cells, each with a central pore (white dots) are clearly visible. (B) Longi-

tudinal section of a gemmae cup shows a lower chamber that produces gemmae and a funnel-shaped upper chamber 

that captures the energy of a falling raindrop and jettisons the gemmae. (C) Still frames from a video of a water drop 

hitting a gemmae cup and transporting gemmae-filled droplets (arrows). Filmed at 3000

 fps

 with a 20-

ms

 exposure.  

Parts B and C are from Edwards et al. (2019). 

PSB 70 (3) 2024

232

Figure 3. Alstroemeria flowers are protandrous, starting as male and switching to female. (A) All six anthers 

are still closed and the style is immature. (B, C) Male phase.  Anthers mature in two groups.  First, three anthers 

curl up and dehisce (B) and then later the remaining three anthers curl upwards and dehisce (C). (D) Female 

phase. The anthers have dropped down and the style has extended, curled upwards and split into three stigma 

lobes each with a pollination droplet.  Stills are from a time-lapse video filmed in the lab over nine days.  The tip 

of the style is indicated by arrows.

capture the energy of a falling raindrop to splash 

and propel mature gemmae, which have risen to 

the surface. Marchantia thus provides a lesson 

in biomechanics and dispersal mechanisms for a 

non-vascular plant. 
If we follow a plant over time, we can document 

movements in plant behavior and gain insight 

in terms of floral design, breeding systems, and 

maintenance of species in nature. Plants are 

unexpectedly agile in their movements that range 

from the explosive flowering in the bunchberry 

dogwood (Cornus canadensis), which opens in 

<0.5 ms (Edwards et al., 2005), to the more subtle 

movements of phototropisms and geotropisms. 

Here I highlight three examples of flowers that use 

movement to switch from one sex to another.  

The first example is Alstroemeria, a genus native 

to South America, but almost always available in 

florist section of local grocery stores. Alstroemeria 

flowers are protandrous, where anthers dehisce 

first; later, the style lengthens, curves upwards, 

and splits into three lobes, each topped with a 

pollination droplet (Figure 3).  Using florists’ 

samples, students can observe these changes 

directly in real time.
The second example is the flower of spring beauty 

(Claytonia caroliniana), which is accessible in New 

England for field observation in the early spring. 

The flowers of Claytonia are also protandrous. 

In a field population, flowers are typically in 

different stages of development. On the first day 

of flowering, stamens are held erect and dehisce 

presenting magenta-colored pollen. On the 

second day, stamens reflex back and the style splits 

into three stigma lobes (Figure 4).

Figure 4

. 

Claytonia caroliniana flowers are protandrous.  On day one of flowering, the stamens are held upright and 

dehisce, presenting pollen. On day two, the stamens bend back toward the petals and the stigma splits into three lobes. 

By day three, most flowers begin to close.

PSB 70 (3) 2024

233

Figure 5. Symplocarpus foetidus flowers are protan-

drous. (A) Spadix with flowers all in female phase. (B) 

Spadix with flowers transitioning with the upper flowers 

in male phase and the lower flowers still in female 

phase. (C) Female phase flowers showing the stigmas 

and styles just protruding from the petals. (D) Male 

phase flowers with stamens, which have extended above 

the petals and dehisced presenting pollen.

The final example is the flower of the iconic skunk 

cabbage (Arisaema triphyllum), which, if available, 

is well worth a field trip to observe the plants in 

situ. This allows one to experience first-hand the 

strong skunky odor, the wet swampy habitat, the 

extraordinary structure of the spathe and spadix, 

and the behavior of the flowers.  Here in New 

England, skunk cabbage is our earliest blooming 

wildflower. It has protogynous flowers (Figure 5) 

but is also amazing in heating up to 35ºC above 

ambient air temperatures with a metabolism 

equivalent to that of a small mammal (Knutson, 

1974), producing a skunk-like odor, and having 

specialized idioblast cells filled with double 

pointed raphide crystals. When broken, idioblasts 

shoot out these glass “spears” presumably as a 

protection against herbivory (see video in Pickett-

Heaps and Pickett-Heaps, 1984). 
Looking closely at plants both in the laboratory 

and in their natural setting can be foundational, 

can serve as a key part of teaching botany, and 

can contribute richly to learning and discovery 

in botany. Most students long remember visiting 

skunk cabbage in its native habitat, or a visit 

to a Sphagnum-dominated kettle-hole bog, or 

even observing the self-digesting flowers of 

Tradescantia, the unfolding and sexual switch 

of an Alstroemeria flower, or the sparkle on a 

Pelargonium petal. The list is endless. By providing 

a full context for plant behavior and enriching 

it with direct observation in both the field and 

the lab, we can give students an entree into new 

discoveries, train their eyes to “see,” and provide 

them the tools to interpret plants and their role 

no matter where they go. If we are to solve our 

twin environmental crises of biodiversity loss and 

climate change, a keen eye and a knowledge of the 

diversity of plants is key. 

REFERENCES

Edwards, J., D. Whitaker, S. Klionsky, and M. J. Las-

kowski. 2005. A record-breaking pollen catapult. Na-

ture 435: 164.

Edwards, J., M. Laskowski, T. I. Baskin, N. Mitchell, and 

B. DeMeo. 2019.  The role of water in fast plant move-

ments. Integrative and Comparative Biology 59: 1525-

1534.

Enquist, B. J., X. Feng, B. Boyle, B. Maitner, E. A. New-

man, P. M. Jørgensen, P. R. Roehrdanz, et al.  2019.  The 

commonness of rarity: Global and future distribution of 

rarity across land plants. Sciene Advances 5: eaaz0414.

Griscom, B. W., J. Adams, P. W. Ellis, R. A. Houghton, 

G. Lomax, D. A. Miteva, W. H. Schlesinger, D. Shoch, et 

al. 2017.  Natural Climate Solutions.  Proceedings of the 

National Academy of Sciences 114: 11645-11650.  

PSB 70 (3) 2024

234

Knutson, R.M. 1974. Heat production and temperature 

regulation in the Eastern skunk cabbage. Science 186: 

746-747.

Lenton, T. M., M. Crouch, M. Johnson, N. Pires, and L. 

Dolan. 2012.  First plants cooled the Ordovician. Na-

ture Geoscience 5: 86-89.

Lenton, T. M., T. W. Dahl, S. J. Daines, B. J. W. Mills, 

K. Ozaki, M. R. Saltzman, and P. Porada. 2016.  Ear-

liest land plants created modern levels of atmospheric 

oxygen. Proceedings of the National Academy of Sciences

113: 9704-9709.

Pickett-Heaps, J., and J. Pickett-Heaps. 1984. 

Living 

Cells: Structure, Function, Diversity [DVD]. Sinauer 

(Oxford University Press).

Pörtner, H.-O., R. J. Scholes, A. Arneth, D. K. A. Barnes, 

M. T. Burrows, S. E. Diamond, C. M. Duarte, et al. 

2023. Overcoming the coupled climate and biodiver-

sity crises and their societal impacts. Science 380: 256.

Qiu, Y. L., Y. Cho, J. C. Cox, and J. D. Palmer. 1999. 

The gain of three mitochondrial introns identifies liv-

erworts as the earliest land plants. Nature 394: 671-674.

Zielinski, C. et al.  2023. Time to treat the climate and 

nature crisis as one indivisible global health  emergen-

cy.  British Medical Journal 383: 2355. [This editorial 

was published in many journals.]

235

Bessey Award Winners Through the Years

By Marshall D. Sundberg

Roe R. Cross Distinguished Professor of 

Biology – Emeritus, Emporia State University, 

Emporia, KS

As a college sophomore in 1968, I was first 

introduced to a novel way of instruction by my 

botany professor, William Muir.  Muir’s approach 

was unique in many ways, starting with the fact 

that he had just lost his sight as a complication 

of diabetes. He lectured, without notes, and 

drew sketches on the board using one hand as a 

placeholder as he sketched—and then quizzed 

us to be sure we understood what was being 

illustrated. If you were the one called on, you 

would have to carefully describe what you saw and 

what it meant—carefully enough that someone 

who could not see it (Dr. Muir) would understand 

what you meant. For the rest of my career, this was 

a tool I would use, particularly in lectures, whether 

it be for small seminars or lectures of more than 

300 students. Several examples are described 

below, and many are also included in Uno et al. 

(2013). (Copies of this book are still available from 

the BSA office: https://crm.botany.org/civicrm/

contribute/transact?reset=1&id=8.)
A second unique approach was to critique the 

textbook, as necessary, during the course of the 

class. This was only done occasionally, and for 

“big” things in the introductory course, but it 

was a main component of upper division courses. 

For the latter, this consisted of mimeographed 

handouts of corrections, elaborations, or current 

research, related to the chapter being discussed. I 

still have many of these as folded chapter inserts in 

undergraduate textbooks I’ve kept in my library.  

I’ll give some examples below of the kind of 

textbook “updating” I used in class. The “mimeo 

Using Inquiry as a Tool to 

Help Students Develop a 

More Sophisticated 

Understanding of Frequently 

Misunderstood Concepts

handouts” remain the model I use when reviewing 

manuscripts and textbooks.  
The third characteristic that set Muir apart was 

his philosophy of science. Virtually every science 

teacher I ever had, including Muir, emphasized the 

power of science in developing an understanding 

of nature. But Muir also emphasized the limits of 

science. The usual way of doing science emphasizes 

finding a solution to a particular problem, but 

this narrow focus often results in unintended 

consequences that might have been avoided if a 

broader perspective was used.  Especially in applied 

science, implementation is often dependent on 

many different non-science constraints: economic, 

legal, environmental, social, religious, and more. 

Finally, the fact that science grows by building on 

the foundation of existing knowledge (accretion) 

makes it very difficult to accept any paradigm-

shifting innovation. I begin with my favorite 

example of a paradigm shift that occurred during 

my career. 

Accepting a Paradigm Shift

Endosymbiotic Origin of Eukaryotic Cells

In the 1960s, it was just becoming accepted 

that bacteria and blue-green algae were closely 

related and shared features termed Prokaryotic. 

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One taxonomic question was “Should at least 

the bacteria be split out of the Plant Kingdom?”  

The author of my textbook (Cronquist, 1961) put 

them together in a single division separate from 

the rest of plants. Regardless, everyone agreed that 

blue-greens evolved from bacteria and the green 

algae probably did as well. Both were the result of 

repeated mutations, recombinations, and natural 

selection over the course of millions of years. At 

the start of my Plant Evolution course, Spring Term 

1970, Muir made us aware that a young biologist, 

Lynn Sagan [Margulis] had published a paper 3 

years earlier suggesting eukaryotic cells arose via 

symbiosis between pre-existing prokaryotic cells. 

Two years later, as a rookie grad student, I listened 

to her plenary address at the 1972 American 

Institute of Biological Sciences Annual Meeting in 

Minneapolis. It was in Northrup Auditorium with 

several thousand biologists present. At the end of 

her talk, half the audience was politely applauding, 

but the rest were jeering! This was my introduction 

to professional scientific meetings and thankfully, 

I’ve never seen anything like it again. Evolution 

by anything other than natural selection was 

considered heretical. Endosymbiosis is one of 

those paradigm shifts that is now well accepted, 

and I’ve told this story every time I’ve taught it. I 

approach this in class by presenting the traditional 

interpretation, the new alternative, then asking 

for what kind of evidence would be necessary to 

support the alternative. Now, here’s the evidence 

and we can move forward.

Primary Root Growth

This is an example where the lecture component 

is covered in “Inquiring about Plants” (pp. 80-85). 

Briefly, I present students with a macrophotograph 

of a growing root tip and ask individuals to 

describe different parts of the image and/or 

speculate on the possible function of a particular 

part. We finally focus on the “naked” tip and switch 

to a photomicrograph of a longitudinal section 

showing the root/root cap junction (Figure 1).
I tell students to make a sketch of the general 

patterns they observe and to predict how cell 

divisions might produce these patterns. Finally, 

based on their interpretation, where would they 

expect most cell divisions to occur? The patterns 

suggest this should be near the arrow in the figure 

and this, in fact, is what was in Cronquist’s (1961) 

textbook. I then show the radiomicrograph and 

explain how it was made (Figure 2). Onion roots 

were grown for 24 hours in a tritiated thymidine 

solution so that any nucleus that underwent 

mitosis would pick up the radioactive tracer. The 

dark spots cover nuclei that picked up the tracer. 

The region we thought would have the highest 

mitotic activity actually has the least: the quiescent 

center.  This was first proposed by Clowes (1950), 

but it was 6 years before he confirmed his theory 

using radioactive tracers as shown above. I then 

challenge the students to devise an experiment 

we could actually do in our laboratory, in a single 

lab period, to confirm the presence of a quiescent 

center. The hint is mitosis and we’ll see this below.
Some other paradigm shifts during my teaching 

career include transposable elements modifying 

the Central Dogma of DNA and the role of 

epigenetics in producing “inheritance of acquired 

characteristics”: a neo-Lamarckian, and even 

neo-Darwinian “gemmules” concept. A possible 

paradigm shift, in process during the last decade, 

relates to consciousness and behavior in plants. 

Schlanger (2024) provides a readable, well-

documented lay account of the current status of 

this theory.

Challenging the Textbook

Primary Growth of Roots

Clowes (1950) first discovered the quiescent 

center by looking at the distribution of mitotic 

figures in longitudinal sections of root tips. This 

is where I lead students in my question above.  

However, what I’m interested in during this lab is 

not just finding evidence of the quiescent center, 

but in examining the relationship between cell 

division and cell enlargement in the growth of the 

root. For the latter, any old onion root tip slide will 

do, but if you also want to identify the quiescent 

center you must use a near-median section. 

When I first developed this activity, I examined 

every onion root tip slide in the department’s 

collection for all courses. Out of more than 200 

slides, only 22 were near median and I set these 

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aside specifically for this lab.  The procedure for 

this lab is detailed in Sundberg (1981). Students 

observe four sequential fields of view, at 40X, 

beginning with the intersection between the root 

cap and root apical meristem at the bottom edge 

of the first field. They must determine the average 

cell length and width from median vertical and 

horizontal files across the field. Estimate the total 

number of cells in the field by dividing the area of 

the field by the area of a single cell and calculate 

the mitotic index (MI) (number of cells showing 

mitotic figures / total number of cells) X 100. 

(An additional benefit of this lab is the necessity 

to do some basic mathematical computations.) 

To indicate the presence of the quiescent center, 

divide the total number of cells in the field by 3 

and separately calculate MI for the estimated 

bottom, middle, and top thirds of the first field of 

view, centered just above the root cap (fields IA, 

IB, and IC). When data collection is completed for 

this field, move the slide so cells at the top edge 

of the original field are now at the bottom of the 

new field of view and observe and collect data 

for the entire field II. This process is repeated for 

fields III and IV.  Plot the data as in Figure 3. In 

general, as you move from the tip to the base of the 

root, the average cell length increases and the MI 

decreases. The low MI in field IA is an indication 

of the quiescent center. These is the types of data 

originally used by Clowes to predict its presence.
Finally, I ask students to make a sketch of the entire 

longitudinal root they observed and then, based 

on their data, label the zones of cell division, cell 

elongation, and cell maturation as is often found 

in textbooks. Figure 4A is from Campbell (Urry 

et al., 2023) and 4B is from Raven and Johnson 

(Raven et al., 2023). Do you see the difference 

in the labelling of these zones?  Which figure is 

supported by the student data in Figure 3? (Hint: 

Are the zones discrete or do they overlap?)

Monocot Stem Structure

One of my favorite examples of challenging 

the textbook in lecture involves the structure 

of monocot stems, and I feature it in “Inquiring 

about Plants.”  Most biology textbooks describe 

Figure 1. Longitudinal section of a maize root at inter-

section between the root cap (below) and tip of the root 

apical meristem.

Figure 2.  Radiomicrograph of an onion root tip in medi-

an longitudinal section. Black covers nuclei that emitted 

radiation by incorporating labelled thymidine into their 

DNA following cell division.

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Figure 3. Comparison of Mitotic Index (open circles) and Average Cell Length (closed circles) along root tip axis from 

junction with root cap (IA) towards root hair region (IV).

Figure 4. Comparison figures of Zones of Cell Division, Elongation, and Differentiation in two popular 

contemporary textbooks. (A) Urey et al., 2023. (B) Mason et al., 2023.

A

B

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the vascular bundles of monocots stems as being 

“scattered throughout the ground tissue” (Urry 

et al., 2023 [p. 768]; Raven et al., 2023 [p. 778]). 

Figure 5A is a photomicrograph of a maize stem 

that I put on the screen for the class. I then ask 

them to observe it carefully and make a sketch, 

filling a full page of their notebook, of the general 

tissue regions they observe. I circulate through 

the class with a blank overhead transparency 

sheet observing the student sketches, but with no 

comments. When I find one that clearly shows 

some patterns, I’ll ask that student to trace her 

sketch on the overhead sheet. Figure 5B is a 

typical example. As a class we’ll then go through 

the sketch noting any observed patterns, labeling 

parts, and adding additional patterns observed by 

other students.
Students often recognize at least six patterns: (1) 

concentric rings of bundles, (2) bundle density 

greater in outside rings than interior ones; (3) 

bundle size greater in interior bundles than 

outside ones, (4) bundles seem to alternate from 

one ring to the next, (5) there is noticeable cell 

differentiation within bundles, and (6) bundles 

always orient in the same direction relative to 

the surface, regardless of where they occur in 

the stem. A close-up photomicrograph (Figure 

6) makes it easier to see cell differences within a 

bundle. A last question, which they’ll turn in on 

a ¼ sheet of scratch paper, is: Which direction is 

the nearest epidermis in Figure 6: left, top, right, 

or bottom?  How do you know? Students identify 

definite patterns of bundles within the stem; 

they are not simply “scattered.” In fact, they are 

precisely arranged, and studies of serial sections 

can predict which bundle of which leaf, up and 

down the stem, every one of these stem bundles 

will supply (Pizzolato and Sundberg, 2002).

Figure 5. (A) Cross-section of maize stem. (B) Student sketch of A.

Figure 6. (A) Magnification of portion of Figure 5A. (B) Student sketch of A.

A

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Plant Migration on Mountains and 

Climate Change

This is another example from “Inquiring about 

Plants” (pp. 135-146). In brief, Humboldt 

suggested that the change in plant communities, 

as elevation increases in the mountains, is similar 

to that observed with increasing latitude on Earth. 

We also know that the combination of average 

temperature and average precipitation in a region 

allows us to predict the plant communities 

(biomes) that will be present. Given the warming 

associated with climate change, what would you 

predict will happen, over time, to the various 

plant communities occurring on the sides of a 

mountain? It seems obvious that as the climate 

warms, plant communities will migrate to higher 

elevations.
The data in Table 1 show the average change in 

elevation for 73 tree species in the Coast Range of 

Northern California since the 1930s. Do the data 

support your prediction for the effect of climate 

change?  Why or why not?  What other factor 

most likely accounts for the unexpected decreased 

elevations in so many species? Hint: go back to the 

two factors we know we can use to predict plant 

communities/biomes we will find in an area.

Size and Distribution of Stomata in 

Desert Plants

My final example is an extension of the stomata 

section of “Inquiring about Plants,” where we 

ask is there a relationship between the number 

of stomata and the environment of the plant (pp. 

45–47)?  It seems logical to predict that there is 

a decrease in stomatal density with increasing 

drought and that stomata should be restricted only 

to the lower surface of leaves in desert plants. One 

of my early students in freshman botany tested 

this for his independent class project and got 

some unexpected results. I followed this up with a 

grant to work at the Desert Botanical Garden near 

Phoenix (Sundberg, 1986). In fact, three-fourths 

of the 111 species examined were amphistomatic, 

and only semi-woody xerophytes had a higher 

frequency on the lower (abaxial) surface than the 

upper surface (Table 2). Leaf and stem succulents 

did have the lowest stomatal densities, but they 

also, unexpectedly, had the largest stomata 

(Figure 7). Some seasonally dehiscent desert trees 

had more than 500 stomata/mm

2. 

In summary, 

classroom inquiry can not only improve students’ 

understanding of the scientific concepts we teach 

but sometimes their naïve, unbiased, observations 

can uncover new connections and expand our 

understanding of science.
Ever since Bill Muir forced me to be an active 

learner through inquiry, I have used this approach 

in my own teaching and learning pedagogy. I 

particularly focus on common misconceptions 

held by many students (and the general public) 

and challenge them with data supporting more 

sophisticated understanding and a philosophy of 

lifetime learning (Sundberg and Moncada, 1994).

Table 1.  Change in elevation of 73 montane tree species 

in the Coast Range of California. Highlighted numbers 

are statistically significant changes. 

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REFERENCES

Clowes, F. A. L. 1950. Root apical meristems of Fagus syl-

vatica. New Phytologist 49: 248-268.

Cronquist, A. 1961. Introductory Botany. New York, Harper 

& Row.

Table 2. Position of stomates on leaves of desert plants.

Figure 7. Trends of stomatal lengths on stomatal den-

sity for various life forms of desert plants: (A) Seasonally 

deciduous; (B) Semiwoody,; (C) Drought deciduous; (D) 

Leaf succulent; (E) Stem succulent; (F) Green stem; (G) 

Evergreen; (H) Ephemeral.  WEISS shows trend of tem-

perate mesophytic plants.

Pizzolato, T. D., and M. D. Sundberg. 2002. Initiation of the 

vascular system in the shoot of Zea  mays  L. (Poaceae) II. 

The procambial leaf traces. International  Journal  of  Plant 

Sciences 162: 353-367.

Raven, P. H., G. B. Johnson, K. A. Mason, J. B. Losos, and 

T. Duncan. 2023. Biology, ed 13. New York, McGraw Hill.

Schlanger, Z. 2024. The Light Eaters: How the Unseen World 

of Plant Intelligence offers a New Understanding of Life on 

Earth. New York, Harper Collins.

Sundberg, M. D. 1981. Making the Most of Onion Root Tip 

Mitosis. The American Biology Teacher 43: 386-388.

Sundberg, M. D. 1986. A comparison of stomatal distribu-

tion and length in succulent and non-succulent desert plants. 

Phytomorphology 36: 53-66.

Sundberg, M. D., and G. J. Moncada. 1994. Creating effec-

tive investigative laboratories for undergraduates. BioSci-

ence 44: 698-704.

Uno, G. E., M. D. Sundberg, and C. A. Hemingway. 2013. 

Inquiring About Plants: A Practical Guide to Engaging Sci-

ence Practices. St. Louis, Botanical Society of America. Fig-

ure 4A is from Campbell (Urry et al., 2023) and 4B is from 

Raven and Johnson (Mason et al., 2023).

Urry, L. A., M. L. Cain, S. A. Wasserman, P. V. Minorsky, 

and J. B. Reece. 2023. Campbell Biology, ed 11.  New York, 

Pearson.

Weiss, A. 1865. Untersuchungen uber die Zahlenund Gross-

verhaltnisse der Spatoffnungen.  Jarbuch fur Wissenschafli-
che Botanik 4: 125-197.

242

Bessey Award Winners Through the Years

How can we encourage our students to look at 

plants like Georgia O’Keeffe did? Slowing down, 

taking time to really look at plants, being a careful 

observer of the living world, appreciating their 

beauty and instilling curiosity to look, notice, and 

go back for more.  
Unfortunately, we learn or are taught impediments 

to learning and curiosity. Do you remember 

the first time you made a drawing of a flower or 

picked a bouquet of dandelions? Did you stop 

drawing because you were not an artist?  Did 

you stop collecting dandelions because they 

are “just weeds”?  We urge teachers of botany to 

both remember our roots, the joy of discovery, 

the historical and contemporary importance of 

drawing in teaching botany, and to further explore 

fine-arts practices outside of traditional botanical 

Don’t Forget Our Roots: 

Learning with Drawing

By Stefanie M. Ickert-Bond

1

and Brett C. Couch

2,3

1 

University of Alaska Museum of the North, 

Herbarium, and Department of Biology and 

Wildlife, University of Alaska Fairbanks, 

1962 Yukon Dr., Fairbanks, AK 99775, USA

2

 University of British Columbia, Depart-

ments of Botany and Zoology, 3156-6270 

University Blvd, Vancouver BC V6T 1Z4, 

Canada

Thesis and Philosophy 

“’If one painted a flower the size it is, nobody would 

look at it. When you take a flower in your hand 

and really look at it,’—and she cupped a strong, 

exquisite hand and held it close to her face—’it’s 

your world for the moment. I want to give that 

world to someone else. Most people in the city rush 

around so they have no time to look at a flower. I 

want them to see it whether they want to or not.’”

        Georgia O’Keeffe 

(in an interview with Mary Braggiotti [1946])

drawing. To rekindle curiosity and the excitement 

of exploration of the botanical world, we propose 

that students should be encouraged to value the 

process not just the product: make it fun.    
The use of drawing, painting, and illustration 

has a long history in botany for a very good 

reason; to draw or paint something, you need 

to look carefully. Although Leonardo Da Vinci 

is well known for his painting, Mona Lisa, he 

also studied human anatomy and botany. His 

approach to science was observational, and he 

filled sketchbooks and journals with detailed 

observations to understand the world he observed 

such as his study of Ornithogalum sp. and other 

plants (Figure 1A). His journals also illustrate how 

he used drawing as part of his thought process as 

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an artist and inventor. In one example he created 

analogies between the leaves of plants and the 

forces of water in the water eddies (Figure 1). 
Our philosophy is that students need to be trained 

to look first and look again, again and again. 

Discover the joy of looking and discovering; 

handling and dissecting plants, exploring and 

documenting local flora, and doing fieldwork 

provides context for other observations. We 

want students to develop observational skills as a 

habit of mind. We want to train students to deal 

with the reality that nature is messy and realize 

that perfect drawings from a textbook rarely 

capture the diversity they will encounter when 

observing plants or other features of the natural 

world. The challenges of observing nature and the 

insights gained from careful observation are also 

highlighted in Da Vinci’s study of moving water. 

He observed and described the three-dimensional 

nature of flowing water, and developed the idea 

that turbulent flows consist of a range of co-

existing eddies, varying in scale from large to small 

(Figure 1B)—but it was not until 1941 that this 

concept was mathematically formalized by A. N. 

Kolmogorov as the “cascade model of turbulence” 

published first in Russian (1941) with an English 

translation not appearing in print until 1991. 

One of our classical mentors is Charles Edwin 

Bessey, who created the first undergraduate 

botanical laboratory in the United States, used 

and encouraged drawing in teaching, and had 

students draw from collected specimens in the 

lab. His motto was “Science with Practice,” and 

he expected students to learn for themselves. A 

quote from his 1896 book The Essentials of Botany

illustrates how he expected drawing to be used as 

part of learning about plants (Figure 2): 

“In the use of this book I must urge that it 

is intended to serve as a guide only to the 

teacher and student. The student must 

actually see as much as possible of what is 

here brought to his notice. The book simply 

marshals in logical order the objects to be 

studied …. the young botanist should not 

be content to obtain all his facts at second 

hand; he must see with his own eyes all that 

may be seen” (Bessey, 1896) 

From here we hope to inspire you to explore 

other approaches to using drawing in your 

classes. 

Figure 1. Drawings by Leonardo Da Vinci. (A) Star of Bethlehem Ornithogalum sp., and other plants c.1506-1512. 

Wikimedia Commons. (B) Studies of water passing obstacles and falling, c. 1508-1509. Wikimedia Commons.

A

B

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How Instructors Use Drawing in 

Teaching 

From our own undergraduate education, we 

enjoyed the instructors who gave us time to 

appreciate botany, those who slowed down 

instruction by using drawings, and those who 

encouraged us to explore the subjects carefully 

and record our observations. Lectures in our 

undergraduate botany courses consisted of the 

professor using vertical-sliding chalkboards 

that were each filled with botanical drawings to 

illustrate the lecture content and were available 

to review after class. Unlike listening to a lecture 

or viewing a static image—activities in which 

students passively absorb information—these 

interactive, progressive, drawings actively engage 

students to record the lecture content. Today, we 

continue this tradition, with some added tech; 

as instructors we make use of drawings in our 

teaching to illustrate plant structures and convey 

information about taxonomically important 

structures. We try to create classes that are both 

engaging and foster slowing down and looking. 

The Learning Glass or Lightboard platform is a 

high-tech version of drawing on the blackboard, 

but with a technological twist. It creates a visual 

connection with the instructor who makes these 

Learning Glass lectures particularly engaging. 

During the use of the Learning Glass, a large piece 

of glass ringed by LED lights, the instructor stands 

behind the glass and uses fluorescent markers to 

draw on the glass, and the ink catches the light 

from the LED and glows clearly. The Learning 

Glass software collapses the perspective of the 

viewer and presenter into one shared perspective, 

allowing students to view the instructor in real 

time drawing and communicating with them, 

while getting visual and textual reinforcement of 

content (e.g., interactive progressive drawing). 

Students presented with classes using the Learning 

Glass had better knowledge retention over the 

same timescale as content delivered through 

PowerPoint (Hennige, 2020). Research has also 

shown that making drawings or sketches increases 

retention of information compared with taking 

in class written notes (Fernandes et al., 2018; 

Higley et al., 2024); in botany, which uses a lot 

of specialized terminology, drawings paired with 

terminology are particularly valuable for helping 

students retain information presented. Drawings 

need not be artistic—instead the drawing process 

is the main educational benefit of drawing, which 

Higley et al. (2024) elaborate on in their “Value 

of Bad Drawing in Teaching.” We recorded many 

Learning Glass lectures for BIOL195 - Introduction 

into Flora of Alaska at the University of Alaska 

(Ickert-Bond and Kaden, 2022) and have made 

these available on Botany Depot (see Appendix) 

and on our class website (https://introtoflora.

community.uaf.edu/module-1/).

Figure 2. Illustration of Bessey’s classification of diatoms 

(Bessey, 1900).

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What Do Students Do: Drawing in 

Labs and in the Field (or “A Pencil Is 

the Best of Eyes”)

The first step in learning to observe is to slow down 

and take time. As Georgia O’Keeffe noted, people, 

like our students, are busy and often don’t take time 

to look.  One approach we have found that sets the 

stage for practicing slowing down and looking is 

to take students out of their comfort zone of the 

science lab and into an art gallery.  Students are led 

through a slow looking activity using the Visual 

Thinking Strategies framework (Yenawine, 2013) 

with artwork that is chosen specifically because it 

is visually complex and something that students 

haven't  seen before. This puts all of the participants 

on the same level in terms of experience with the 

work and so they cannot easily draw on previous 

knowledge or preconceptions. Students spend 

approximately 30 minutes looking at a work and 

responding to the prompts: (1) What do you 

see? (2) What makes you say that? and (3) What 

else do you see? This activity sets the tone for 

the entire semester in the lab. Students are then 

introduced to a variety of drawing and sketching 

activities that are typical in studio arts classes and 

that are intended to practice observing rather 

than producing finished drawings. An example 

is gesture drawing (Figure 3A–B). This is a very 

fast, timed, drawing of a subject—typically 15 to 

60 seconds.  Students are given various objects 

(pinecones, fern leaves, flowers) and given only 15 

to 60 seconds to quickly capture as much of the 

object as possible. The drawings often look like 

scribbles; the purpose is not to capture a realistic 

representation, but rather to practice seeing the 

entire thing and recording some general features 

or ideas. A second type of activity is blind contour 

drawing, a slow looking activity. The idea here is to 

take a longer period (10–30 minutes) and slowly 

“trace” the contours (edges of a subject). The 

observer needs to convince themselves that the 

pencil is actually touching the edge of the object 

as they slowly move the pencil over the paper to 

draw the contour (Nicolaïdes, 1975). The catch 

here is that the student is not to look at the paper 

while they are drawing. The entire focus is on the 

contour of the object. By design, the drawing 

will not be a perfectly accurate representation of 

the object being drawn—the drawings are often 

rather funny—but the purpose is not a completed 

drawing, but to practice focused observation. 

Through the term, students will also practice 

the manual dexterity skills required for making 

drawings by doing simple doodles and activities 

that focus on making different types of marks. 

Each lab period begins with a doodle activity 

and some activity that focuses on some specific 

element of observation—layered drawings to show 

movement or sequential observations (Figure 

3D). (Couch et al., 2023).  Students then apply 

these skills to making sketches of microscopic 

structures or organisms. 
We see that the skills of summarizing, simplifying, 

and observing, practiced with various drawing 

activities, provide an inroad to further 

development of visual literacy in students. For 

Figure 3. Examples of different drawing and sketching 

techniques by Brett Couch (2023). (A) Gesture draw-

ing of dandelion, (B) details added to part A, (C) detail 

drawing of leaf venation, and (D) layered drawing of 

amoeba. 

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246

example, the ability to interpret and comprehend 

visual information in the sciences like graphs, 

figures, models, and diagrams increased by: (1) 

using sketches to develop or communicate ideas, 

and thinking through problems like diagramming 

an experimental design, or making predictions 

about patterns of data consistent with a particular 

hypothesis; (2) using visual media to communicate 

effectively in the form of figures or graphical 

abstracts; and (3) providing a mechanism of 

visualizing abstract concepts such as a gene on a 

chromosome.  

Multiple authors have recognized the value 

of drawing across biology and STEM for 

communication and learning (Waldrip et al., 

2010; Ainsworth et al., 2011; Landin, 2011; Tyler 

et al., 2018). Landin (2013) summarized the 

importance of drawing: 

“It’s weird how much 

visual information I miss until I draw an object. 

Our brain just skips over details that don’t fit with 

our preconceptions. When we draw, we have to 

include everything—and that leads to learning.”

We encourage you to think about ways of creating 

experiences for your students that engage them to 

use drawing iteratively and repeatedly, and in ways 

that promote curiosity, thinking, and learning—

to reveal the joy in slowing down and making a 

flower, a leaf, or a whole plant their world for a 

moment. 

ACKNOWLEDGMENTS

The authors acknowledge the generous support 

of a CITE fellowship (Chancellor’s Innovation 

in Teaching & eLearning Program, UAF) to 

S.I.B. for developing the asynchronous BIOL190 

Introduction to Alaska Flora course and support 

from eCampus, and the amazing instructional 

designers, especially Christen Booth, eCampus 

Creative Director, as well as support for Brandon 

Corder, and Nkosi Evans (the University of 

Wisconsin, Madison) and Todd Widhelm (The 

Field Museum of Natural History) from the 

United States National Science Foundation 

(DBI-2001509) for the collaborative project, 

Building a Global Consortium of Bryophytes and 

Lichens: Keystones of Cryptobiotic Communities 

(GLOBAL; https://globaltcn.utk.edu/) to support 

the creation of the lichen and bryophyte learning 

glass videos at UAF. Thanks also to Dr. Shelly 

Rosenblum (Curator of Academic Programs 

at the Morris and Helen Belkin Gallery, UBC), 

Holly Schmidt (artist and educator), and the 

Morris and Helen Belkin Gallery and for their 

work in developing and hosting activities for 

biology students at UBC. We also like to thank 

the organizer of the symposium at Botany 2024, 

“Bessey’s Legacy: Enthusiasm and Innovation 

in Botanical Instruction,” for supporting our 

participation and encouraging this submission 

and the many mentors and teachers who have 

shared their passion for teaching and drawing.

REFERENCES

Ainsworth, S., V. Prain, and R. Tytler. 2011. Drawing 

to learn in science. Science: 333: 1096–1097. 
Bessey, C. E. 1896. Essentials of Botany, ed 8. New 

York: Henry and Holt Company. 
Bessey, C. E. 1900. The modern conception of the 

structure and classification of diatoms, with a revision 

of the tribes and a rearrangement of the North Ameri-

can genera. Transactions of the American Microscopi-

cal Society 21: 61–86. 
Braggiotti, M. 1946. Her Worlds Are Many. New York 

Post (16 May 1946), 45.
Couch, B. C., H. Schmidt, and C. Goedhart. 2023. 

Training a biologist’s mind through an artist’s eye. Ad-

vances in Biology Laboratory Instruction 43: 1–15.
Fernandes, M. A., J. D. Wammes, and M. E. Meade. 

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Appendix

I. Talk at the Bessey Symposium at Botany 

2024 in Grand Rapids, Michigan

• The Google Slides can be found 

at:

https://docs.google.com/presentation/

d/10YNrZVBC0pA2JLgRLYG-

cumqrZ5lVEIZsAUqt1ME7uU/

e

II. Learning Glass Lectures (LGLs)

• Most LGLs can be found on the BIOL190- 

Introduction to Alaska Flora website, 

under the individual modules, here are 

those for module 1: https://introtoflora.

community.uaf.edu/module-1/

• Angiosperm life cycle: https://media.uaf.

edu/media/t/0_lxrrkn0o

• A complete listing of LGLs can be found 

on Botany Depot https://botanydepot.

com/2020/03/13/online-course-intro-

to-alaska-flora-by-stefanie-ickert-bond/

Four new LGLs were completed in spring 2024:

• Bryophytes Versus Lichens Comparison 

https://media.uaf.edu/media/

t/1_0k9hwi0o

• How to ID Mosses - https://media.uaf.

edu/media/t/1_d75j0o1k

• Life Cycles of Bryophytes and Lichens 

https://media.uaf.edu/media/t/1_

b7j7ba5i

• Basic Lichen Biology - https://media.

uaf.edu/media/t/1_avubjnps

III. Virtual Herbarium

• https://www.thinglink.com/

scene/1406090479749038081  

248

Bessey Award Winners Through the Years

By Bruce Kirchhoff

Emeritus and Adjunct Professor of 

Biology, University of North Carolina, 

Greensboro, NC

There are two aspects to great teaching: The first 

and most important is to be yourself and to share 

yourself with your students in ways that enhance 

their learning. The second is to make sure that 

your students do most of the work. Learning these 

lessons took me almost 25 years of classroom 

experience and resulted in several major teaching 

awards, including the Bessey Award. I share my 

insights here in the hope that it will not take you 

quite as long for similar achievements.
The best way to share yourself in the classroom is 

to present with enthusiasm. Let your students see 

your love for your subject. Your enthusiasm will 

reach them better than any content you deliver.  

It sounds easy, but presenting with enthusiasm 

without losing intellectual focus takes practice. 

One thing that helped me to show my enthusiasm 

was to begin each lecture with a joke related to my 

course content. Since, at the end of my career, I 

most frequently taught Plant Diversity and Plant 

Systematics, all of my jokes were related to these 

subjects. What worked best for me was to find 

a visual joke related to the course content and 

present it at the beginning of class, just before I 

asked opening student-response questions. If I 

could not find a joke that fit the class, I created 

one. The jokes I chose were never wildly funny, 

but they were entertaining. For instance, I found 

this joke on the web: There is a picture of an 

abandoned car that is covered with ivy with the 

question, “Why are plants capable of consuming 

cars?” I would enthusiastically ask the class “Well, 

why?” After several wrong guesses, I would reveal 

The Two Rules of Great 

Teaching: Present with 

Enthusiasm and Make Your 

Students Do the Work

the answer on the next slide: “Because they are 

auto-trophic.” Occasionally a student would get 

this correct and I would react with joy, throwing 

my arms up and almost shouting “YES!” and 

maybe adding “Someone was paying attention in 

Intro Bio!” 
I would often continue this light-hearted teasing 

during other parts of the class. For instance, my 

quizzes and exams always included a question 

on mitosis and miosis. Although I did not cover 

these subjects in my classes, I felt that the students 

should all have a basic understanding of the 

difference between mitosis and meiosis before 

they graduated. For some students these questions 

provided free points, but more than half the class 

regularly missed them. Right at the beginning 

of the semester, during the first lecture, I would 

say something like this: “You all learned about 

mitosis and meiosis in introductory biology. 

Some of you have had genetics, where you learned 

about them again. How many of you remember 

the difference?” (Maybe 2 people out of 24 raise 

their hands.) “That is what I thought [said with 

great humor]! Well, in this class you will have the 

opportunity to test your knowledge because every 

quiz, every test, and many clicker quizzes will 

PSB 70 (3) 2024

249

have a question about mitosis and meiosis. For 

some of you these will be free points [said with 

enthusiasm]! That is great! I want to give you free 

points! However, for some of you these questions 

will be a source of never-ending frustration [said 

with a hint of sadness]. You will always be asking 

yourself, ‘How did I miss that AGAIN?!’ Take my 

advice. Review the difference between mitosis and 

meiosis so you are not pulling your hair out at 

the end of each quiz [I mime pulling out my hair 

and show them my bald head]. You see where this 

leads!” By miming pulling out my hair, I show the 

students that I am just as foolish as they are—that I 

have made all the same mistakes. This, and similar 

gestures throughout the class, gives the students 

permission to accept their own mistakes with 

grace, and even to laugh at them. I have found that 

this attitude does much more to enhance learning 

than any serious admonitions I might use.
Later in the class when I gave a clicker question 

on mitosis or meiosis and 60% of the class got it 

correct, I would enthusiastically congratulate them 

and then speak to the 40% of the class that missed 

the question. I might again mime pulling out my 

hair and remind them that these are supposed to 

be free points and that if they do not want to end 

up like me (I tap my bald head) then they should 

really review mitosis and meiosis. Then after class 

I might post some links to good review sites. Some 

students will continue to miss these questions no 

matter what I do, but even if they never learn 

the difference between mitosis and meiosis, the 

class atmosphere that I create with these types of 

interactions helps the students feel comfortable 

and encourages them to work hard.
Let me give another example of how I used 

enthusiasm and jokes to promote student learning. 

One fall break I went to the beach for a few days. 

While there I drew one of the life cycles we were 

learning in the sand and took a picture of it. That 

picture was the opening slide of the first lecture 

after break. The caption read “Why? What do 

you do at the beach?” (Figure 1). Although most 

of my students worked at least half time and had 

been working over break, this joke reminded 

them that they should not forget the class material 

just because they were otherwise engaged. Jokes 

like this reminded them of the seriousness of the 

material without hitting them over the head with it.
Although there are many other ways to present 

with enthusiasm, I hope that these examples give 

you some idea of how I approached my class. You 

can see that there is a definite advantage to being 

bald. 
This brings me to my second point: Make the 

students do the work. Presenting with enthusiasm 

creates a class atmosphere that is conducive to 

learning, but the students must still learn the 

material. The only way to do this is for them to do 

the hard work of learning. I believe that our task 

as instructors is to make this hard work seem like 

fun, at least as much as possible. I do not mean 

to minimize the work that the students have to 

do. I strongly believe that the only way to learn is 

through hard work. However, if the students will 

not do the work you assign, they cannot learn. An 

example of what could be an extremely effective 

learning method will make this clear. I call this the 

White Paper Method. 
The White Paper Method starts with a sheet of 

blank paper, a pencil, and a copy of the material 

that the student wants to learn. This can be their 

lecture notes, notes provided by their instructor, 

or their textbook. Once the student has identified 

the material for their study session, they close 

their notes and take out the paper and pencil. They 

Figure 1. A life cycle drawn in the sand.

PSB 70 (3) 2024

250

then recreate, with as much detail as possible, the 

material they want to learn. Let’s say that they 

want to learn the life cycle of a vascular plant like 

Pinus. On their blank piece of paper, they draw the 

life cycle in as much detail as they can. It does not 

matter if they can draw only a small portion of the 

life cycle. Once they have done this as best they can, 

they compare their work to their notes and correct 

their work so that they have a perfectly drawn life 

cycle. It is best if they do this in a different color 

so they reinforce what they have yet to learn. Now 

comes the most important part. They take their 

corrected work and throw it away and take out a 

blank piece of paper. On this paper they draw the 

life cycle again. Since they just reviewed it, they 

will do a better job. When they correct this version 

as they did the first, there will be fewer corrections. 

If they have not gotten it fully correct, they repeat 

the White Paper process until they can draw the 

life cycle perfectly from memory. That ends this 

study session for this content. If they repeat this 

process at least one more time before the exam, 

they will ace any questions on this life cycle. The 

White Paper Method is extremely effective! I have 

had a student go from failing at midterm to a B 

in the class by using this method. That means she 

went from failing every test, to getting an A on 

every test. This is an amazing accomplishment.
The problem with the White Paper Method is 

that the students will not do it. The student I just 

mentioned was a soccer player and if she had failed 

my class she would have been kicked off the team. 

She was successful because of this incentive and 

because her coach made her use the White Paper 

Method. Most students do not have this incentive 

and, for whatever reason, will not use this method.
This is our conundrum as teachers. We must find a 

way to get the students to do the work that they are 

unable to do on their own. There are several ways 

I approached this problem. Perhaps some of them 

will appeal to you.
One of the most powerful ways of promoting 

student learning is to create effective homework. 

The archetype of effective homework is the White 

Paper Method, but we already know that students 

will not do this on their own. Can we fool them 

into doing it with creative assignments? One way 

I found to do this involves weekly in-class quizzes. 

In my Plant Diversity class, I expected the students 

to be able to draw even complex life cycles from 

memory. To get them to do this, I would first 

draw the life cycle with them in class, sometimes 

asking them to draw it from memory during the 

class period and then going over the life cycle with 

them as they corrected it on their papers. You will 

recognize this as the first iteration of the White 

Paper Method. To get them to continue the process 

at home, I would tell them which life cycles were 

candidates for next week’s quiz. Early in the class 

there were few choices, but late in the semester 

there were so many that telling them that any life 

cycle was fair game would overwhelm them and 

they would do poorly. If I told them to expect one 

of the following three life cycles on next week’s 

quiz, they would make sure that they could draw 

them from memory and the vast majority of the 

students would get full credit. This made grading 

very simple. I only needed to glance at a life cycle 

to see that it was correct. Grading could be done 

quite quickly. Over the course of a few weeks, I 

could cover all of the required life cycles with 

near-perfect performance. In this case I used in-

class quizzes to create the incentive for students to 

do the work on their own.
As every teacher knows, one only really learns 

the material when one has to teach it to others. 

With this in mind, I began requiring my students 

to present some of the lectures in my classes. In 

Plant Systematics, the students presented almost 

all of the plant families. I presented a few at the 

beginning of the class to give them examples of 

what I expected, then the students took over, 

presenting the family characteristics for the rest of 

the semester. I gave them very explicit instructions 

on what to include (see links below), and most 

students did very well. To my surprise they did 

even better when they presented online during 

Covid. I suspect that this was because I allowed 

them to present with their cameras off, which 

relieved much of their anxiety. 
When I taught Biological Evolution, I presented 

the first few lectures before turning the rest of the 

material over to the students. My approach to this 

PSB 70 (3) 2024

251

material was unusual in that I had the student’s 

present chapters from The Origin

(1

st 

ed),

and 

a graduate text by Kemp, Fossils and Evolution. 

I found that Darwin spoke to the students in 

ways that contemporary texts could not. Darwin 

was writing to an audience that doubted the 

correctness of his theory. He wrote to persuade, 

not to present facts. This approach was much 

more meaningful to my students, many of whom 

came from religious backgrounds where they were 

encouraged to avoid any discussion of evolution. 

A fuller description of my approach in this class 

can be found at the following link: https://sites.

google.com/view/active-learning-in-use/ .
Another great way to get students involved in 

a class is to have them take notes. For years I 

wondered how I could do this without requiring 

the students to turn in their notes, which would 

require an inordinate amount of grading. The 

Covid pandemic provided an opportunity to 

try a new method, with good success. I was 

teaching Plant Systematics when my university 

closed. Since the students presented many of the 

lectures in this class, I had to use synchronous 

class time for these lectures. To accommodate 

this, I decided to record my lectures and present 

them asynchronously. In order to ensure that the 

students were paying attention, I required them to 

turn in their notes using our course management 

system (Canvas). Most students took notes on 

paper and photographed them using one of the 

many phone apps created for this purpose. Many 

of my students used Genius Scan. I used a three-

tier grading system for these notes: good (100%), 

needs work (50%), no credit (0%). After a few 

lectures I could show the students examples of 

good notes (with the note-taker’s permission). 

Soon 98% of the students were getting full credit. 

This made grading very easy. In fact, I spent more 

time waiting for the digital files to load in Canvas 

than I did grading the student’s work. Some of 

the notes were amazingly good. I still wonder 

if it would be possible to get students to take 

good notes in face-to-face lectures, but I never 

succeeded in this before I retired. Perhaps some 

variation on these procedures will work for you.

In closing, it would be remiss of me to fail to 

mention my work creating visual learning 

software. I will not go into detail about this 

software here because it is described more fully 

elsewhere (see links below). The software is free 

and Open Source. I tested it in the classroom and 

found it to be extremely effective (Kirchoff et al., 

2014). Stephanie Jeffries at North Carolina State 

has created an online version and extension of this 

software for use in teaching plant identification. 

The links to her work are also below.

• Active Visual Learning Software: https://

sites.google.com/view/image-quiz/home

• Teaching materials for a course on Plant 

Diversity: https://osf.io/69skm/

• Plant Life Cycle Diagrams: 

http://planted.botany.org/index.

php?P=FullRecord&ID=578

• Recorded lectures on Plant 

Diversity: https://www.youtube.com/@&lt;/p>

plantdiversity

• Recorded lectures on Plant Systematics: https://

www.youtube.com/@Plant_Systematics

• White Paper Method: https://youtu.be/

Gyu4KQPekx0

• Stephanie Jeffries ILEX (Identify-Learn-

Explore) online tool: https://sites.google.

com/ncsu.edu/ncstatedendrology/ilex-study-

tool?pli=1

REFERENCES

Kirchoff, B. K., P. F. Delaney, M. Horton, and R. Del-

linger-Johnston. 2014. Optimizing Learning of Scien-

tific Category Knowledge in the Classroom: The Case 

of  Plant  Identification.  CBE-Life  Sciences  Education

13: 425-436.

252

Bessey Award Winners Through the Years

Growing up in Detroit and attending public 

schools, I thought I would become a doctor, lawyer, 

or writer. At that time, I had never heard of doing 

science for a career. But from my first Biology 

lab course at the University of Michigan, going 

on a walk outside looking at trees and insects, I 

discovered the great outdoors. In the end, I chose 

graduate school to let me continue doing just that. 

Undergrad Days

My Botany education was fun and solid, with most 

courses delivered in the standard lecture format. 

The professors were talented lecturers, enthralling 

us with subject matter and amusing us with their 

personalities (Botany Professors Hiroshi Ikuma, 

Herb Wagner, Ed Voss; Zoology Professor Dan 

Janzen; Organic Chem with Professor Richard 

Lawton). I was a very good student and excellent 

note-taker, recording everything they said almost 

verbatim. Writing it all down helped me commit 

it to memory. Reading textbooks and papers, 

solving problems, and reviewing notes helped 

me succeed in almost every course. Some courses 

were less conventional. Dan Janzen’s Habitats 

and Organisms course consisted of non-stop 

lectures, to a huge audience in a darkened room, 

while showing us beautiful slideshows of animals 

and plants from around the world.  In John 

Vandermeer’s Quantitative Ecology, we served 

as guinea pigs for his early textbook/workbook 

of problems (Vandermeer, 1981).  Biochemistry 

used the self-directed “Keller Plan,” taking tests 

on every chapter complemented with lectures on 

extra material. Field experiences in courses at the 

University of Michigan Biological Station and as a 

research assistant to Sally Kleinfeldt in the woods 

The Evolution of an Educator

By Suzanne Koptur

Professor Emerita, Florida International 

University

of New Hampshire let me see what research might 

be like. Although I also worked as a nurse’s aide for 

two summers (one in Detroit at Plymouth General 

Hospital, the other at Mott’s Children’s Hospital), 

which gave me a view of the medical world, I chose 

the path that would be more fun, with perhaps less 

job security and money, but more time outdoors 

and doing things I loved. 
I had work-study employment in the University 

Herbarium, working with Dr. Robert Shaffer to 

index the type collection of Fungi. I got some 

research experience with Dr. Rogers McVaugh, 

writing a Latin description of a newly discovered 

species of Pedicularis from Mexico (my first 

publication: McVaugh and Koptur, 1978). I 

wrote a senior thesis about extrafloral nectaries 

with a focus on aspen under the advice of Dan 

Janzen. During my undergrad time, I was lucky 

to be an assistant to Teaching Fellows (TFs as 

they were called, and we were TAs) in Practical 

Botany (taught at the Botanical Garden) and 

Plant Systematics on campus. Though headed 

in the Systematics direction, I shifted to Ecology 

because it seemed there was an endless supply of 

PSB 70 (3) 2024

253

interesting questions to investigate. I received a 

Noble Fellowship from the Smithsonian Tropical 

Research Institute, doing my first tropical field 

work on Barro Colorado Island. 

Graduate Studies in California: A 

Whole New Flora! (And Then Some!)

Entering grad school at Berkeley in the Botany 

Department as Teaching Assistants (TAs) in 

General Biology, we were required to take a 

weekly teaching seminar in addition to our twice-

weekly TA meetings for General Biology, a two-

quarter sequence directed by Dr. Bill Jensen. 

In the TA meetings, we learned the content and 

how to run the lab sessions, but in the seminar, 

we learned about good practices in science 

education. In my group of four TAs who all taught 

labs at the same time, two were grad students in 

the SESAME program (Search for Excellence in 

Science Education), which held a great appeal for 

me. Both suggested it might be best to stay in pure 

science, since I could always move to their field of 

science education later if I chose, but moving in 

the other direction might be harder. 
After Gen Bio, I was a TA (with many others) for 

California Flora with Robert Ornduff, and a new 

basic botany course with Don Kaplan (along with 

fellow grad student Darlene DeMason). At Berkeley 

we taught labs, but we were also required to attend 

the corresponding lectures. I moonlighted as a 

note taker for the lecture courses in which I taught 

labs, for Black Lightning, a service run from a copy 

store where notes were copied and made available 

to subscribing students. I remember writing and 

typing them on mimeo sheets! I was unaware that 

this practice was controversial, as it evolved into 

some professors selling their lecture notes, etc. 

Nowadays (in fact, within the following decade), it 

is more common practice to provide lecture notes 

as part of the course materials for many professors.
I took some wonderful courses as a graduate 

student at Berkeley, including Evolutionary 

Ecology taught by Herbert Baker, who highly 

valued teaching as a pursuit. In that course there 

were students from many departments (Botany, 

Forestry, Zoology, Entomology), providing 

connections for all of us with other parts of the 

university. Herbert’s lectures and demonstration 

labs were full of information, letting us take as much 

time as we wanted with his collection of articles, 

books, plant and animal examples, artifacts, etc. 

That was also the way he taught Economic Botany 

(see Baker, 1978).  We solved problems using basic 

statistics and were expected to do a project of our 

own design. The field experiment I carried out at 

Tilden Park (taking the bus up into the Berkeley 

hills each time) was material for my first solo 

publication (Koptur, 1979). I learned a lot about 

prioritizing projects and publishing from James 

Hickman (thanks, Jim!), who had recently come 

to Berkeley with his wife, Carol Hickman. I also 

got to take the Organization for Tropical Studies’ 

Tropical Biology course in 1977, a full immersion 

introduction to the neotropics, doing faculty-

led group and student-initiated field projects for 

several months.
I then spent 2.5 years away from teaching as RA on 

a grant received by my major professor, Herbert 

Baker, and entomologist Gordon Frankie, to study 

Phenology and Pollination in the Costa Rican 

Cloud Forest, a wonderful time of my life (Koptur 

et al., 1988). After the field work was done, I 

returned to Berkeley to write up my dissertation 

on the plant/animal interactions of Inga and was 

again a TA, for Plant Systematics, and then was an 

RA in the University herbarium. My final semester 

I was asked to teach the Plant Ecology lecture and 

lab because the regular professor (Rob Robichaux) 

was on sabbatical. He graciously shared all his 

notes with me, and I got my first insight into 

preparing for lectures in the traditional way. With 

Suzanne Morse as my TA, we had an adventurous 

and very fun semester with lots of field trips and 

field exercises in interesting places.

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254

Postdoctoral Work in the Midwest 

and Across the Pond

Completing my Doctorate in spring, and after 

unsuccessfully applying for many faculty positions 

in 1982, I took a teaching postdoc at the University 

of Iowa with Hank Howe, where I taught General 

Zoology labs and was in the company of some 

great tropical ecologists. This provided support 

(both financial and intellectual) while I wrote and 

published papers from my dissertation work and 

helped me get a NATO postdoc where I worked 

with John Lawton at the University of York, 

another wonderful time of my life. During these 

postdoc and early faculty times, I was also an 

investigator in the Naturalist-Ecologist Training 

Program during several summers at the University 

of Michigan Biological Station, a great experience 

for mentoring undergrads in independent research 

projects while pursuing my own research. I co-

coordinated the Organization for Tropical Studies 

course in the summer of 1985 with my old buddy 

from undergrad days at the Michigan Biological 

Station, Bob Marquis—my first year as an assistant 

professor (see below).

Training Paid Off

All that teaching experience helped me get a job 

as an assistant professor at a young state university 

in Florida, Florida International University (FIU), 

where I was hired as a population biologist in 

1985. I initially taught Gen Bio 2 and a graduate 

course in Evolutionary Ecology, as FIU was 

working toward an independent MS program. I 

later taught Introduction to Biological Research 

in our new graduate program and started teaching 

Ecology, then Plant Ecology in alternate years, 

along with workshops in Pollination and Field 

Techniques in Plant/Insect Interactions. I also 

stepped into Introductory Botany when David Lee 

and Jenny Richards moved on to other courses. I 

got some great ideas from workshops I took at the 

Botany meetings, including using portfolios in 

non-majors’ courses (thanks, Joe Armstrong!) and 

great hands-on materials for groups, passing out 

sections of the same stem or tree branch (thanks, 

Stokes Baker!). I also participated in Project 

FIRST—Faculty Institutes for Reforming Science 

Teaching Through Field Stations—and was part 

of the FIU team for several iterations of this NSF-

funded project. I had previously arranged field 

trips to Archbold Bio Station (Swain, 2019) for 

some of my courses, but this brought together a 

community of educators from different institutions 

in Florida to learn new approaches and design 

activities that could be used by all, especially in 

the field. 

Time for a Change

After 10 or more years of delivering material in 

lectures, I was getting a little bored teaching in 

the same old ways. Enticed by a workshop at the 

annual Ecological Society meeting, I learned about 

Innovative Methods in Large Lecture Courses 

from two inspiring scientists and educators: Diane 

Ebert-May and Carol Brewer. That workshop 

was really life-changing for me! I learned how to 

foster more interactions with students and among 

students (Ebert-May et al., 1997). I realized that 

average attention spans are short, so that after 12–

15 minutes of listening, most students were zoning 

out. By introducing active learning breaks that 

broke up the twice-weekly 75-minute classes into 

three or more sections, the students were engaged 

and got to talk and/or move around, breathing 

new energy into the lecture hall. I accepted the 

challenge and encouragement to transform my 

lecture courses, but one lecture/day at a time, and 

over several offerings of each course. 
I was teaching Ecology every year, and so I took 

their advice with transforming this required 

course for all Biology majors. For three years I did 

an experiment with a different topic to see if active 

learning made a difference. The topics used in this 

experiment were Energy in Ecosystems (Spring 

2006), Biological Communities (Spring 2008), 

and Adaptation and Natural Selection (Fall 2009). 

The content was not assumed to be comparable 

PSB 70 (3) 2024

255

among the topics, but each served as a vehicle for 

the experiment.  I had an ideal set-up for teaching 

the same material two ways, as half the class went 

to receive instruction from our science librarian 

about finding articles in the scientific literature, 

while the other half attended an Ecology lecture. I 

taught the same topic twice each time, but in two 

different ways.
To test the hypothesis that active student 

engagement results in greater learning, I used the 

following experimental design.  All students in 

the experiment were to read the same textbook 

chapter and view the same material in the lecture 

that I delivered (i.e., the same PowerPoint slides), 

and each would write an in-class essay (“minute 

paper”).  Students in the Active Learning group 

would have three in-class active-learning breaks 

during the lecture, e.g. a “think/pair/share,” 

making a categorizing grid, concept map making, 

class modeling, human tableau, etc.—ideas I got 

from a great resource, my favorite education 

“cookbook,” Classroom Assessment Techniques 

by Angelo and Cross (1993). 
I could see which students attended the library 

session on each day, and which ones were present 

in lecture on the other day via the in-class essays 

they turned in. By using data from relevant 

questions on a pretest, from material on the topic 

in the mid-term exam, and then in the final exam, 

we saw that students who participated in the 

Active Learning version of the topic learned more 

and demonstrated this by better performance on 

the relevant questions on the mid-term exams 

(X1) than those students taught in the traditional 

way (Figure 1). There were greater gains for 

students in the Active lecture than for those in 

the regular lecture in the mid-term results (QX1 

vs. Qpre). This difference did not hold up long 

term: performance on the final exam questions, 

QF vs. Qpre, did not differ significantly between 

those two groups. An interesting side result was 

that students who attended either type of lecture 

showed greater gains (by every measure except the 

mid-term exam) than those who did not attend 

lecture! I presented these results in a poster at a 

Gordon conference (organized by Gordon Uno) 

at Bates College, where it was energizing and 

inspiring to meet with science educators from all 

over. I continued to change my lectures in this 

class, and in others, over the following years.  

Changes at the University 

FIU created a STEM Transformation Institute 

(https://stem.fiu.edu/), in which I was one of a 

group of founding faculty fellows. We participated 

in many workshops on teaching and learning, 

assessment, and different ways of engaging and 

inspiring students. I learned more about active 

learning methods, starting with lab activities—

presenting students with challenges and some 

materials, then letting them explore to answer 

problems. I learned about professors who 

simply did not lecture in class, rather using the 

lecture time to have students work together and 

Figure 1. Summary of outcomes for students attending 

traditional lecture (vertical striped bars) versus Active 

Learning lecture (diagonal striped bars) and those who did 

not attend lecture (empty gray bars).  On the x-axis, Final 

= score on final exam, Pretest – score on Pretest, X1 = score 

on midterm exam in which the topic was covered, QX1 = 

score on questions on the topic on the mid-term exam (a 

specific part of X1), Qpre = score on questions on the topic 

on the pretest, and QF = score on questions on the topic 

on the final exam. Total = total score in course. Data are 

combined for three different topics in three different trials 

(semesters of the course), so normalized gain makes the re-

sults comparable among trials.

PSB 70 (3) 2024

256

independently on thinking and problem solving. 

A Learning Assistant Program was started, and 

grew, employing undergraduate students who had 

previously taken a course to help the professor 

manage group work in larger courses. We also 

adopted Peer-Led Teaching and Learning (PLTL) 

in many of the required majors’ courses, where 

current students attend a weekly session led by a 

student who has previously taken and done well in 

the course.  This intervention helped students do 

better than those who did not have PLTL as data 

from my Ecology courses show (Figure 2). Exam 

averages were higher for students taking PLTL 

along with the lecture course than those who did 

not (78.5 vs. 73.3), and substantially more passed 

the course as well (78.6% vs. 62.3). PLTL can help 

students from marginalized groups succeed in 

STEM majors (Sloane et al., 2021). 
The Stem Transformation Institute also developed 

a Discipline Based Education Research group 

(DBER) that holds seminars biweekly throughout 

the semester—a great chance to learn from 

outside experts and others at FIU, and to interact 

with faculty in one’s own and other departments. 

It connected those of us teaching science classes 

with science and math education colleagues and 

fostered collaborations, leading to many projects 

and publications. DBER meetings were enjoyable, 

and it was great to get to know others across the 

university who cared about teaching despite more 

praise for research accomplishments.  

Flipping Lecture: Fun and Beneficial 

for Students and Faculty

I decided to flip my courses, and this was an 

exciting time for me and the students (although 

some pre-med students worried they would not 

learn enough in my classes). Each class meeting 

had active learning almost all the time, working in 

groups with the supervision and help of Learning 

Assistants (LAs), presenting to others, and using 

white boards and other means of communication. 

Students were to prepare for each class by reading 

the assigned textbook chapter, listening to a 

couple of short PowerPoint lectures I had pre-

recorded, and checking out (reading or listening 

to) other resources I posted online, and taking a 

quiz over the textbook chapter contents. We used 

clickers in class to provide instant feedback on 

multiple-choice questions, and then had students 

discuss questions and answer again. I started 

giving assessments (exams) in class using a two-

part system: first, students would take the exam 

individually and turn in their answer sheets; then 

with their group they would work through the test 

and fill out IFAT (Instant Feedback Assessment 

Test) bubble cards. Each person’s score was the 

average of the two. Attendance was very good 

because we always had activities and most of them 

“counted” as part of the students’ grades. 
Comparing student performance in my Ecology 

courses over all the years I taught the traditional way 

(lecture with no LAs) with the flipped course with 

LAs (taught only in the spring semesters) shows 

a marked improvement in course completion, 

passing rate (Figure 3A), and distribution of final 

course grades (Figure 3B).  However, comparing 

the performance of students in two summer 

semesters of online Ecology (synchronous), one 

with and one without LAs, showed no difference 

(Figure 4); in fact, there were more high achievers 

without LAs. 

Figure 2. The difference Peer-Led Teaching and Learning 

(PLTL) made in Ecology: participation of Ecology stu-

dents in PLTL vs. Final course grade earned. PLTL yes 

= students who attended PLTL sessions; PLTL no = stu-

dents who did not attend PLTL.

PSB 70 (3) 2024

257

Pre-Adaptation for Remote Teaching 

When the pandemic struck and we switched 

to synchronous online teaching via Zoom, this 

flipped teaching format was easy to adapt, because 

lectures and quizzes were already online and the 

students were expected to go through the online 

materials and take the chapter quizzes before class. 

On Zoom, groups could work in breakout rooms 

with each LA visiting a subset of the groups. The 

groups (or a selected few of them) would then 

summarize or present to the entire class. 
To meet community demands and enhance 

financial gain for the university, the College of 

Arts and Sciences wanted to develop a Biology BA 

degree fully online. Because I had taught Ecology 

A

Figure 3. Comparisons of the Ecology course taught 

flipped (with Learning Assistants, LA) versus traditional 

lecture (No LA).  (A) Passing rates: multiple offerings 

over the years combined by semester, sample sizes for 

each given in the yellow bars; (B) Grade distributions in 

the same courses, including drops (DRs), sample sizes as 

in (A).

Figure 4. Comparisons of the online synchronous Ecolo-

gy course taught in flipped style with Learning Assistants 

(LA) versus more traditional lecture style without activi-

ties (No LA) over summer term offerings (initial summer 

class size was 50; the next time it was 97). (A) Pass/fail 

comparisons; sample sizes for each given in the yellow 

bars; (B) Grade distributions, including drops (DRs).

A

B

B

PSB 70 (3) 2024

258

many times, I volunteered to develop Ecology for 

this curriculum as asynchronous online.  Each 

topic was covered by short lectures, activities to 

be done by one student, quizzes on each chapter, 

and exams that could be taken twice. I developed 

an online lab, using some SimBio resources and 

creating one third of the labs myself with help 

from the teaching assistants (thank you especially 

to Cleo Pimienta and Andrea Salas Primoli!). This 

course has become increasingly popular over time, 

as have other online offerings of required courses, 

allowing working students and parents of small 

children to do this on their own schedule. 

Community Education

Along with university activities, I have always been 

willing to give talks and organize activities for the 

public by lecturing and holding workshops for 

plant societies, nature groups, elementary, middle, 

and high schools. Some examples include the 

“After School Gardening Gang” with elementary 

students, project PRIDE (Pine Rocklands in 

Dade Environments) at West Miami Middle 

School (teacher Lisette Perez Munoz received 

a Toyota Tapestry grant), and several projects 

with the environmental magnet at TERRA 

Environmental Research Institute. All have helped 

me to communicate better, learning how to reach 

students of different abilities at all levels. Working 

in groups, the students help and teach each other, 

with more positive results for all. 
All of these changes have certainly transformed 

my teaching and students’ learning over time. 

Most of the changes were gradual (breaking 

up the lecture with activities), but some were 

extreme (flipping the lectures in all my courses).  

This is kind of how evolution proceeds: gradual 

versus punctuated equilibria. I have always liked 

teaching and consider it an important part of a 

professor’s job, even though FIU became more 

and more research and funding oriented over my 

decades there. Evolving my teaching by adapting 

my methods to a changing clientele has helped me 

retain my interest in and enthusiasm for teaching 

for over 40 years. 

REFERENCES

Angelo, T. A., and K. P. Cross. 1993). Classroom As-

sessment Techniques: A Handbook for College Teach-

ers, ed 2. San Francisco: Jossey-Bass. 
Baker, H. G. 1978. Plants and Civilization, ed 3. Wad-

sworth Publishing.
Ebert-May, D., C. A. Brewer, and S. Allred. 1997. In-

novation in Large Lectures: Teaching for Active Learn-

ing. Bioscience 47: 601-607. 
Koptur, S. 1979.  Facultative mutualism between 

weedy vetches bearing extrafloral nectaries and weedy 

ants in California.  American  Journal  of  Botany 66: 

1016–1020.
Koptur, S., W. A. Haber, G. W. Frankie and H. G. Bak-

er. 1988.  Phenological studies of shrub and treelet spe-

cies in tropical cloud forests of Costa Rica.  Journal of 

Tropical Ecology 4: 323346.
McVaugh, R., & S. Koptur. 1978. A new species of 

Pedicularis from Jalisco, Mexico. Contr. Univ. Mich. 

Herbarium 11(5): 298-300.
Sloane, J. D., R. D. P. Dunk, J. J. Snyder, C. I. Win-

terton, K. M. Schmid, and J. R. Wiles. 2021. Peer-

Led Team Learning is Associated with an Increased 

Retention Rate for STEM Majors from Marginalized 

Groups. Proceedings of the 13th Annual Research 

Symposium, National Association of Biology Teachers 

2021, pp. 1-9.
Swain, H. M. 2019. All of life’s experiences count at a 

biological field station. Frontiers in Ecology and Evo-

lution 17: 102-103. 
Vandermeer, J. 1981. Elementary Mathematical Ecol-

ogy. Wiley, New York. 

259

From the PSB Special Issue on Art in the Botanical Sciences

Within the past year, the Plant Science Bulletin has published two special issues in the special anthology, 

Art and the Botanical Sciences: Past, Present, and Future (the Fall 2023 and Spring 2024 issues). These 

issues grew out of our first workshop on botanical art at Botany 2022 in Anchorage, AK, and the collected 

articles explored many facets of the importance of botanical arts. 
We present two more articles that were unable to appear in those issues: “Illustrating Cretaceous Park: 

First steps toward a botanical field guide for the Hell Creek Formation” by Kirk R. Johnson and Marjorie 

Leggitt as well as “Reconstructing the botanical past: Art and paleobotany” by Edward J. Spagnuolo et 

al. We hope you enjoy these articles and encourage you explore the past special issues at https://botany.

org/psbarchive/view/issues!

The SciArt Collective

Nicolette Sipperly, Stony Brook University • Rosemary Glos, University of Michigan

Kasey Pham, University of Florida • Patricia Chan, University of Wisconsin-Madison

Ashley Hamersma, University of Florida

SPECIAL SECTION 

Art in the Botanical Sciences: 

Past, Present, and Future

260

From the PSB Special Issue on Art in the Botanical Sciences

Illustrating Cretaceous Park: First 

Steps Toward a Botanical Field 

Guide for the Hell Creek Formation

By Kirk R. Johnson

1,  

and

Marjorie Leggitt

2

1

National Museum of Natural History, Wash-

ington, D.C. 

2

Boulder, CO 

[All renderings and models 

©Marjorie Leggitt]

ABSTRACT

Fossil plants provide unique data that can 

lead to credible reconstructions of ancient 

terrestrial landscapes and ecosystems. This paper 

describes our process as we use art and science 

to reconstitute the vegetation of the last North 

American dinosaurs (with apologies to extant 

birds). Our art-science toolkit includes geology, 

sedimentology, palynology, precision excavation 

and censuses of fossil plant sites, accurate tracing of 

fossil leaves and flowers, comparative analysis with 

modern plant relatives, articulated reconstruction 

drawings of fossil material, construction of 

schematics showing floral architecture and 

phyllotaxy, application of traditional and not-so-

traditional artistic methods, and the completion of 

botanical image plates. Scientifically accurate plant 

species portraits are then combined with similarly 

generated animal reconstructions, and geologically 

constrained topography and geomorphology to 

create plausible views of lost worlds.

The Dinosaur Renaissance began in the late 1960s 

with John Ostrom’s discovery of Deinonychus, a 

wolf-sized predatory dinosaur with claws on both 

hands and feet, and Bob Bakker’s lively renderings 

of agile and active dinosaurs. When Stephen 

Spielberg’s Jurassic Park movie debuted in 1993, 

Ostrom’s dinosaur was labeled Velociraptor and 

the film portrayed terrifyingly realistic animals. 

Paleoart had become “pop art,” but there were 

other problems too. The paleobotanist played 

by Laura Dern complained that the protagonists 

needed the opinion of a paleobotanist, and she 

was right. While the dinosaurs of Jurassic Park 

were largely from the Cretaceous Period, the 

surrounding vegetation was simply that of modern 

Hawaii. 
At the same time, as an artist-paleobotanist team, 

we were working on actual fossils from the Hell 

Creek Formation of North Dakota to reconstruct 

a true Cretaceous Park. The resulting diorama in 

the Prehistoric Journey exhibition that opened in 

1995 at the Denver Museum of Nature & Science 

included a walk-through forest foliated with more 

than 24,000 plastic leaves, all of them based on 

actual fossil leaves (Johnson, 1996; Leggitt and 

Johnson, 1999). Never had a dinosaur diorama 

been vegetated with plants that were collected 

in direct association with the dinosaurs. The ten 

plant species we reconstructed for this diorama 

have gone on to be the plant palette for the Late 

Cretaceous and have been featured in many 

PSB 70 (3) 2024

261

subsequent paintings, books, cartoons, dioramas, 

and video games. Continued excavation over the 

last 30 years has yielded a remarkably diverse Hell 

Creek flora with more than 300 species (Johnson, 

2002) 
We are now embarking on an effort to bring 

botanical reality to the vegetation that was the base 

of the food chain that produced Tyrannosaurus 

rex, the planet’s greatest terrestrial apex predator. 

We plan to do this by focusing on a suite of the 

best Hell Creek Formation fossil leaf quarries 

that we have collected over the last 30 years. 

These quarries represent different stratigraphic 

levels on the 100-m-thick formation and different 

depositional settings including ponds, floodplains, 

riverbeds, and levees. These quarries were chosen 

because they have superb preservation, commonly 

yield complete leaves, and show high plant 

diversity. Each quarry will yield the data needed to 

reconstruct a specific time and place from the last 

1.5 million years of the Cretaceous. In this article, 

we demonstrate how we reconstruct a single plant, 

Cobbania hickeyi, using an example from the 

“Licking Leaves” site, a pond deposit in Harding 

County, northwestern South Dakota (Denver 

Museum of Nature & Science locality 2703). 

Materials and Methods

Leaves and other plant parts are typically buried 

in clay, mud, or sand in or near rivers and ponds 

and are preserved as compressions or impressions 

in claystone, mudstone, or sandstone. Subsequent 

uplift and erosion create the outcrops that are 

the source of fossil plants. During fossilization, 

original leaf organic matter is typically degraded 

or destroyed, leaving a leaf-shaped void in the 

rock. This fact is useful because the rock will break 

along this plane of weakness to yield imprints of 

both the top and the bottom of the leaf. 
During fieldwork in 1994 in Dinosaur Provincial 

Park, Alberta, and in southwestern North Dakota, 

we collected two separate examples of a complete 

floating aquatic plant with a rosette of leaves that 

we interpreted to be inflated. Stockey et al. (2007) 

described this plant, named it Cobbania corrugata, 

and assigned it to an aquatic clade of the Araceae. 

In this paper, we reconstruct the closely related 

species,  Cobbania hickeyi (Stockey et al., 2016), 

which was based on one complete plant and 

many loose leaves from the Licking Leaves quarry 

(Figure 1). 
Close collaboration between artist and scientist is 

extremely important throughout the illustration 

process, and for this plant we relied on our 

colleagues Ruth Stockey and Gar Rothwell. 

Reconstructing a three-dimensional plant from 

a flattened and sediment filled fossil required 

both mental and physical models. To do this, the 

artist (M.L.) traced several leaf fossils, “restoring” 

each in its entirety, and paying close attention to 

shapes, margin, and venation (Figure 2A). She 

used the drawings to create and arrange paper and 

wire leaf models to view the plant from various 

perspectives. 

Figure 1. A single leaf Cobbania hickeyi as it was found 

in the Licking Leaves quarry. The inflated part of the leaf 

has lifted off to show the interior venation of the leaf.

PSB 70 (3) 2024

262

The resulting top-down linear schematic illustrated 

the plant’s spiral phyllotaxy, proper leaf size, and 

arrangement (Figure 2B). Using this sketch and 

referencing live specimens of Pistia corrugata, she 

fleshed out a detailed rendering of a plant rosette 

from above showing five leaves and a new leaf bud 

(Figure 2C).  
A low-angle photo of the top view rotated the plant 

from a top view to a 3/4 view (Figure 2D), and the 

resulting image portrays the altered shapes and 

position of leaves in relationship to one another at 

an oblique angle (Figure 2E). Because the Cobbania

leaves were inflated in life, it was useful to create a 

clay model to understand how they would appear 

while floating in water (Figure 2F). The clay 

model provides a physical form that facilitated the 

drawing of the leaves and petioles, both above and 

below an imaginary waterline. Shining a light on 

the clay models allowed the creation of a realistic 

interpretation on light on form (Figure 2G). The 

final drawing was transferred to watercolor paper 

where the artist used a living relative, Limnobium, 

for color reference to complete the painting 

(Figure 3).

The dinosaurs of the Hell Creek Formation are 

surely the most illustrated animals of all prehistory. 

It is our goal to reconstruct the vegetation of their 

world with precision and beauty, one species at a 

time (Figure 4).

Figure 2. (A) Tracing and restoration of leaf fossil. (B) Linear schematic of spiral phyllotaxy. (C) Delineated render-

ing of rosette. (D) Low-angle photo of inked illustration. (E) Pencil sketch of plant in oblique angle perspective. (F) 

Clay model helps to “see” how light falls on 3D leaves. (G) Value drawing with highlights and shadows to show form.

Figure 3. Full-color reconstruction of the Cobbania 

hickeyi floating rosette. 

PSB 70 (3) 2024

263

Figure 4. Pencil study for a rendering of a Late Creta-

ceous pond environment with rosettes of Cobbania hick-

eyi floating in a shallow pond covered with of Brasenia 

(watershield).

REFERENCES

Johnson, K. R., 1996, Description of seven common 

fossil leaf species from the Hell Creek Formation 

(Late Cretaceous: Upper Maastrichtian), North Da-

kota, South Dakota, and Montana. Proceedings of 

the Denver Museum of Natural History, series 3, v. 

3, p. 1-48.
Johnson,  K.  R.  2002.  The  megaflora  of  the  Hell 

Creek and lower Fort Union Formations in the 

western Dakotas: vegetational response to climate 

change, the Cretaceous-Tertiary boundary event, 

and rapid marine transgression. In: J. Hartman, K. 

R. Johnson, D. J. Nichols (eds). The Hell Creek For-

mation and the Cretaceous-Tertiary Boundary in the 

northern Great Plains: an integrated continental re-

cord of the end of the Cretaceous. Spec Pap 361, pp. 

329–392. Geological Society of America, Boulder, 

CO.
Leggitt, M. C., and K. R. Johnson. 1999. From fossil 

plants to scientifically-accurate dioramas: the fabri-

cation of prehistoric ecosystems. Journal of Natural 

Science Illustration 3: 3-6.
Stockey, R. A., G. W. Rothwell, and K. R. Johnson. 

2007. Cobbania corrugata gen. et comb. nov. (Ara-

ceae):  a  floating  aquatic  monocot  from  the  Upper 

Cretaceous of western North America. American 

Journal of Botany 94: 609–624.
Stockey, R. A., G. W. Rothwell, and K. R. John-

son. 2016. Evaluating relationships among floating 

aquatic monocots: A new species of Cobbania (Ara-

ceae) from the Upper Maastrichtian of South Da-

kota. International Journal of Plant Sciences 177: 

706–725.

264

From the PSB Special Issue on Art in the Botanical Sciences

By Edward J. Spagnuolo

1,5

, 

L. Alejandro Giraldo

1

, Mario 

Coiro

2,3

, and Susannah Lydon

4

1

Department  of  Geosciences  and  Earth 

and  Environmental  Systems  Institute, 

Pennsylvania  State  University,  University 

Park, PA, USA.

2

Department of Paleontology, University of 

Vienna, Vienna, Austria.

3

 Ronin Institute for Independent Scholarship, 

Montclair, NJ, USA. 

4

School  of  Biosciences,  University  of 

Nottingham, Loughborough, UK.

5

Author  for  correspondence  (email: 

spagnuolo@psu.edu)

ABSTRACT

Paleoart is an important tool for paleobotanists 

when reconstructing fossil plants and ancient 

ecosystems, and communicating with diverse 

audiences. Plants are fundamental components of 

terrestrial ecosystems. Thus, accurately depicting 

ancient plants in art is crucial for communicating 

comprehensive knowledge about ancient life. Here, 

we briefly review the history of paleobotanical 

art, discuss the challenges when accurately 

depicting plants in paleoreconstructions, and 

highlight recent works that reconcile isolated 

plant organs into scientifically accurate whole-

plant and landscape-level reconstructions. 

Historically, paleoart has included plants as 

Reconstructing the Botanical Past: 

Art and Paleobotany

background elements in art featuring charismatic 

vertebrates, resulting in poorly depicted plants and 

ecosystems. Plant blindness—the phenomenon 

in which humans are more inclined to detect 

and appreciate fauna than flora—is a persistent 

problem for science communicators, botanists, 

and paleobotanists. Although plant blindness 

is rampant in 20th-century paleoart, modern 

paleoart that accurately incorporates and focuses 

on ancient plants can increase plant visibility in 

portrayals of the geologic past. 

KEYWORDS

art, fossils, paleoart, paleobotany, plant awareness 

disparity, plant blindness, plant fossils,

scientific reconstructions 

Art is an important tool for scientists to engage 

with both scientific and general audiences (Lesen 

et al., 2016). Paleontological art—or paleoart—

has been used to reconstruct extinct organisms 

and environments for almost 200 years and 

has influenced many of our assumptions about 

the past (Davidson, 2008; Stroud, 2008; Witton 

et al., 2014; Clary et al., 2022b; Manucci and 

Romano, 2022). Paleoart can also be useful to 

better understand and advance paleontological 

paradigms—most famously, the extensive 

updated paleoart that accompanied the Dinosaur 

Renaissance of the late 20th century (McDermott, 

2020). Paleoart includes drawings and paintings, 

museum reconstructions and sculptures, as well 

as documentaries, movies, and even video games; 

here, we will mostly reference drawings and 

paintings, the most common form of paleoart.

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Plants are fundamental for ecosystems and 

society, supporting biodiversity, terrestrial 

biomass, ecosystem structure, and as critical 

food and oxygen sources for humans and other 

organisms. Unfortunately, general audiences, 

policymakers, and other scientists are more likely 

to recognize and appreciate animals compared 

to plants. This disparity, termed plant blindness 

(also known as plant awareness disparity in recent 

years) has been attributed to reduced funding 

for plant-related projects compared to animal-

focused research, as well as a global decrease in 

plant-centered education, conservation, and 

recognition (Wandersee and Schussler, 1999; Drea, 

2011; Balding and Williams, 2016; Jose et al., 2019; 

Margulies et al., 2019; Parsley, 2020; Brownlee et 

al., 2021; Stagg and Dillon, 2022; Stroud et al., 

2022; Walton et al., 2023). 
Paleontology is widely thought of as a “gateway 

science” to other fields in science, technology, 

engineering, and mathematics (STEM), and as a 

way to teach broader audiences larger scientific 

concepts such as evolution, mass extinctions, 

climate change, and biodiversity (Moran et al., 

2015). Often, these education and outreach 

initiatives include, or center on, paleoart (Burns 

et al., 2003; Clary et al., 2022a; Lipps et al., 2022). 

Additionally, plant fossils show how environments 

have responded to climate change, and knowledge 

of fossil history can be used as a rationale for 

the direct conservation of plants and ecosystems 

(e.g., the UNESCO World Heritage Gondwana 

Rainforests of Australia; Young and McDonald, 

1987; Burnham, 2001; Wilson et al., 2011; Ivory et 

al., 2016; Lézine et al., 2019; Kooyman et al., 2020). 

Accurately representing fossil plants in paleoart is 

fundamental for conveying information about life 

in the past.
Paleoart has tended to focus on animals, with 

plants seen as a backdrop or scene-setting, rather 

than as “central characters” (however, see Benca 

et al., 2014; Sanders, 2014; Beans, 2022; Benca, 

2022). Here, we discuss how plants have been 

depicted in paleoreconstructions over time within 

the context of plant blindness. We also consider 

the challenges facing plant paleoart and present 

promising trends for the future. 

BRIEF HISTORY OF PLANT 

PALEOART

Duria antiquior (“A More Ancient Dorset”), 

painted by Sir Henry Thomas De la Beche in 1830 

(Figure 1A), is widely considered the first example 

of a new genre of art: the reconstruction of life in 

the past based on scientific evidence (Rudwick, 

1992, 2014; Lescaze, 2017). Although largely 

a marine scene, this first paleoreconstruction 

included palms and other less easily identifiable 

vegetation on background landmasses. In the 

lithograph versions, produced from De la Beche’s 

work by George Scharf, fern-like and cycad-

looking plants are also recognizable (Rudwick, 

1992; Sharpe, 2022; Sharpe and Clary, 2022).
The circulation of lithographic prints of 

Duria antiquior began the proliferation of 

paleoreconstructions as a means of conveying 

information about life in the deep past to broad, 

non-scientific audiences from the 1830s onwards 

(Clary et al., 2022a), and these illustrations 

frequently incorporated detailed plant 

reconstructions (Vujaković, 2019; Manucci and 

Romano, 2022). Christian Hohe’s final lithograph 

for Georg August Goldfuss’ Petrefacta Germaniae, 

produced in 1844, is an exquisitely detailed scene 

from the Coal Measures with a key detailing the 

plant taxa, demonstrating that Goldfuss expected 

his audience to be as interested in them as in 

animal fossils (Rudwick, 1992).
The importance and ubiquity of coal in people’s 

everyday lives (Yuval-Naeh, 2019), combined 

with popular interest in ferns and their allies 

(Whittingham, 2012), meant that paleoart 

focusing on Carboniferous plants was widespread 

in the latter half of the 19th century (Figure 1B). 

For instance, Carboniferous plants featured in 

Franz Unger’s Die Urwelt in ihren verschiedenen 

Bildungsperioden (“The Primeval World in Various 

Developmental Periods”) published in 1851, 

with artwork by Josef Kuwasseg, which inspired 

Edouard Riou’s illustrations for Louis Figuier’s 

La terre avant le deluge (“The Earth Before the 

Flood”) in 1863 (Rudwick, 1992; Davidson, 2015; 

Vujaković, 2019; Collins, 2022).

Figure 1. Representative examples of plant paleoart throughout history and modern plant-centered paleoart. (A) Henry De 

la Beche’s Duria antiquior. Note palms on the middle-right and some less easily identifiable vegetation on the middle-left. (B) 

Lycophyte, sphenophyte, and pteridosperm taxa from the Carboniferous of the United States depicted in Underwood (1896; 

artist unknown), in turn based on Dana (1874). (C) Dinosaur-centered reconstruction of the Late Cretaceous of Argentina, 

with some minor plant elements in the back (Araucaria) and front right (Zamuneria) (artist: Jorge Antonio González, 

modified from Paulina-Carabajal et al., 2021). (D) Dinosaur-centered reconstruction of the Late Cretaceous of Canada, 

with more prominent plant elements covering the ground (ferns), background (conifers), and with which the dinosaurs are 

interacting (angiosperms) (artist: Julius T. Csotonyi, modified from Mallon and Anderson, 2013). (E) Paleoenvironmental 

reconstruction of the Late Cretaceous of Argentina based on pollen data, which provides a more regional signature. Plants 

depicted include ferns, palms, and conifers (artist: F. Guillén, modified from Barreda et al., 2012). (F) Paleoenvironmen-

tal reconstruction of the mid-Cretaceous of West Antarctica based on pollen, geochemical, sedimentological, and organic 

biomarker data, providing a more accurate depiction of the landscape. Plants depicted included Cyathea (Cyatheaceae), 

Podocarpaceae, and Araucariaceae (artist: James McKay, modified from Klages et al., 2020). (G) Fossil material and recon-

struction of the Early Cretaceous conifer Krassilovia mongolica and the associated leaf morphotaxon Podozamites harrisii. 

From left to right: Articulated seed cones, leaves, winged seeds; and reconstruction of a branch of K. mongolica reconciling 

all of the fossil elements including alternately arranged P. harrisii leafy shoots (artist: Pollyanna von Knorring, modified 

from Herrera et al., 2020). 

All images used here are either Public Domain or have full CC-BY 4.0 rights (https://creativecommons.org/licenses/by/4.0/). 

(A) Duria Antiquior [https://commons.wikimedia.org/wiki/File:Duria_Antiquior.jpg] by Henry De la Beche, 1830. Pub-

lic Domain (B) Carboniferous Pteridophyta [https://commons.wikimedia.org/wiki/File:Our_Native_Ferns_-_Carbonifer-

ous_Pteridophyta.jpg#filelinks] by Lucien Marcus Underwood, 1896. Public Domain. (C) © 2021 Paulina-Carabajal et al., 

CC-BY-4.0 (Paulina-Carabajal et al., 2021). (D) © 2013 Mallon, Anderson, CC-BY-4.0 (Mallon and Anderson, 2013). (E) 

© 2012 Barreda et al, CC-BY-4.0 (Barreda et al., 2012). (F) © 2020 Klages et al., CC-BY-4.0 [https://www.nature.com/

articles/s41586-020-2148-5/figures/3] (Klages et al., 2020) (G) © 2020 Herrera et al, CC-BY-4.0 (Herrera et al., 2020).

PSB 70 (3) 2024

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The “Classic era of paleoart” began in the 1890s in 

the United States with the hugely influential work 

of Charles R. Knight (Milner, 2012; Witton, 2018). 

Knight was famously commissioned to create 

paintings and murals for some of the largest natural 

history museums in the United States (including 

the American Museum of Natural History and the 

Field Museum). Often collaborating extensively 

with vertebrate paleontologists, Knight’s murals 

centered on the charismatic extinct vertebrates 

at the forefront of paleontological discovery with 

naturalistic, but often homogenous, vegetation 

(Vujaković, 2019). However, Knight conducted 

detailed research on the Gilboa forests of New York 

and communicated with paleobotanist Winifred 

Goldring to maximize the paleobotanical accuracy 

of his plant-centered mural Devonian Forest (on 

display at the Field Museum; VanAller Hernick, 

2003). Meanwhile, in Europe, Czech painter 

Zdeněk Burian painted lavish reconstructions 

including flora from Devonian to Quaternary 

times (Lavas, 2016; Witton, 2018).
Unfortunately, the paleoart of the mid-late 20th 

century pushed plants into the background. 

Dinosaurs and other charismatic vertebrates were 

the centerpieces of most paleoart from this time, 

and plants were rarely given much consideration. 

Monkey puzzle trees (Araucaria), cycads 

(Cycadales),  Williamsonia (Bennettitales), palms 

(Arecaceae), and tree ferns (e.g., Cyatheales)—a 

very small fraction of the known fossil floral 

diversity—made up the majority of paleoartistic 

reconstructions of Mesozoic vegetation. The 

majority of known Mesozoic seed plants were 

rarely featured in dinosaur habitats and museum 

reconstructions of the time (Philippe et al., 2009; 

Sanisidro and Barrón, 2016; Herrera et al., 2020). 

Dinosaurs were often reconstructed standing on 

dry, lifeless earth with a handful of nondescript 

monkey puzzle trees in the distance, a plant-blind 

art style coined by Kirk Johnson as “monkey 

puzzles and parking lots” (Johnson and Troll, 

2007; Figure 1C). 

The rise of the Internet and digital art at the 

end of the 20th century enabled a paleoart 

community to develop and thrive online (Witton, 

2018). Although tetrapod-centered approaches 

continued to dominate paleoart at the start of the 

21st century (Figure 1D), some artists deliberately 

flipped this orthodoxy, such as Robert Nicholls 

in his reconstruction of the early Cretaceous 

Antarctic Peninsula (McKie, 2011), and influential 

practitioners such as Witton (2018) have 

advocated for far greater consideration of plants 

by paleoartists (Figure 1E–G).

CHALLENGES TO PLANT 

PALEOART AND THE 

POTENTIAL FOR SPECULATION

The fundamental challenge in paleobotany 

and plant paleoart is creating whole-organism 

reconstructions (Martine et al., 2019) given the 

fragmentary nature of the plant fossil record 

(Spicer and Thomas, 1986). The shedding and 

differential preservation of various plant organs—

including leaves, wood, cones, flowers, spores or 

pollen, as well as fruits and seeds—throughout 

the plant life cycle result in a multitude of 

disarticulated fossils produced by the same 

plant (Dilcher, 1974; Kvaček, 2008; Wilf, 2008a; 

Manchester et al., 2014; Cleal et al., 2021), and 

whole-plant preservation is exceedingly rare 

(e.g., Boucher et al., 2003; Zamaloa et al., 2006). 

Additionally, these isolated fossil organs are often 

named as separate species (or even genera), which 

can be confusing for non-experts and paleoartists. 

For example, a single Carboniferous lycopsid tree 

could be the source of at least six separate fossil 

species if found in isolation (Spicer and Thomas, 

1986). Similarly, the use of morphotaxa—species 

or genera representing a certain morphology 

rather than a biological unit—can be confusing 

for paleoartists (Figure 1G). For example, the 

wood genus Araucarioxylon and the leaf genus 

Brachyphyllum were produced by multiple conifer 

groups (Philippe et al., 2009; Philippe, 2011) but 

are often reconstructed as Araucaria, fueling their 

overuse in paleoart. 

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Although leaves are the most abundant plant 

macrofossils, leaf morphology can be highly 

variable and plastic, even on leaves of the same 

plant; most paleobotanists today use caution 

when taxonomically identifying isolated fossil 

leaves (Dilcher, 1974; Doyle, 2007; Wilf, 2008a; 

Spagnuolo et al., 2022). During the 19th and 20th 

centuries, numerous angiosperm leaves from 

the Cretaceous and Cenozoic were inaccurately 

assigned to extant genera and families, largely 

due to superficial similarities. This has led many 

paleoartists, especially during the 20th century, 

to include genera that were likely not present 

(such as Quercus, Populus, Acer, and Salix) in 

late Cretaceous and early Paleogene landscape 

reconstructions. Although reproductive organs—

such as fruits, seeds, flowers, and cones—are the 

basis for most modern fossil plant taxonomy and 

identification, they are often more delicate and 

produced at much lower abundances than leaves 

(Gastaldo, 1992; Cleal et al., 2021).
When reconstructing ancient ecosystems, 

paleoartists must also consider the scale at which 

they are working. Compressed leaves have been 

shown to mostly represent a snapshot of local 

vegetation, with low levels of non-local influences 

(Burnham, 1994, 1997; Wing and DiMichele, 1995; 

Cleal et al., 2021). Conversely, pollen and spore 

data can represent regional vegetation from many 

habitats within a larger region (Behrensmeyer et 

al., 2000; Birks et al., 2016). When combined, these 

data can be used to accurately depict local (e.g., 

beside a pond) to regional (basin-level) vegetation 

(Figure 1E and F; Opluštil et al., 2014; Costamagna 

et al., 2018; Barreda et al., 2020; Wilf et al., 2022). 

When depicting ancient landscapes, paleoartists 

should also consult with scientists from other 

geological disciplines (e.g., sedimentologists) to 

understand the paleo-topography of the region 

and how that would influence the distribution of 

past vegetation. 
While paleobotany deals with fragmentary 

evidence, illustrations often require a well-

developed organismal concept, often based on 

comparative morphology or nearest living relative 

approaches (Witmer, 1995; Witton, 2018; Martine 

et al., 2019). The nature of the plant fossil record 

and the difficulties associated with reconstructing 

whole plants (Bateman and Hilton, 2009) imply 

a certain degree of speculation regarding the 

reconstruction of most plant fossils. Although 

the practice of representing “known unknowns” 

has become an important part of vertebrate 

paleoart (Conway et al., 2013; Nieuwland, 2020), 

paleoartists seem to be more cautious with plant 

reconstructions.
The reason for such caution could be a lack of 

accessibility to botanical and paleobotanical 

knowledge, as well as limited input from scientists. 

Since the late 19th century, paleoart has been 

driven by commissions, most often by vertebrate 

paleontologists, not paleobotanists. Scientists 

must provide artists with more paleobotanical 

information when possible; however, this can be 

a challenge because plants and animals require 

different environmental settings to fossilize 

and often are not found in the same rocks 

(Behrensmeyer et al., 2000). Navigating the jargon-

rich botanical and paleobotanical literature can 

be incredibly difficult for non-experts, especially 

given the decrease in botanical education in 

general curricula over time (Drea, 2011; Stroud 

et al., 2022). Although botanical illustration is a 

well-established field with a rich history spanning 

centuries (Ben-Ari, 1999; Swann and Pye, 2019; 

Bienvenue and Chare, 2022), paleoartists rarely 

come from a formal background in botanical 

illustration (Sutton, 2019; Dart and Coiro, 2022; 

von Knorring and Coiro, 2022) and instead have 

more varied professional stories (Orr, 2019). 

The expansion of paleoart-focused education in 

traditional botanical illustration curricula might 

provide a way forward to better integrate these 

two fields. 

THE FUTURE IS BRIGHT FOR 

PLANT PALEOART

Over the last 20 years, scientists have made 

massive advancements in understanding plant 

evolution and ancient ecosystems due to the 

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advent of molecular data, mass digitization of 

natural history collections, and new imaging and 

statistical methods (Donoghue and Doyle, 2000; 

Bebber et al., 2010; Amborella Genome Project, 

2013; Page et al., 2015; Coiro et al., 2019; Leebens-

Mack et al., 2019; Bakker et al., 2020; Hedrick 

et al., 2020; Romero et al., 2020; Johnson et al., 

2023). Plant paleoart has also made significant 

strides in accurately reconstructing ancient plants 

and paleo-landscapes (see art in Phillips and 

DiMichele, 1992; DiMichele et al., 2007; Benca et 

al., 2014; Hetherington et al., 2016; McElwain et al., 

2021; Beans, 2022; Benca, 2022). Fossil discoveries 

worldwide have yielded additional fossil plants 

with connected organs, allowing for more accurate 

whole-plant artistic reconstructions (art in Sun 

et al., 1998, 2002; Hermsen et al., 2009; Zhang et 

al., 2010; Opluštil et al., 2014; Gomez et al., 2015; 

Bodnar and Escapa, 2016; Rothwell et al., 2022). 

Extinct plant lineages, which often lack whole-

organismal concepts, are being reconstructed 

and properly included in landscapes (Philippe et 

al., 2009; Barreda et al., 2012; Wang et al., 2012a; 

Herrera et al., 2020). Cretaceous charcoalified 

flowers, and their incredibly detailed artistic 

reconstructions by Pollyanna von Knorring and 

others, have provided an unexpected window into 

early angiosperm evolution (Crepet et al., 2004; 

Schönenberger, 2005; Crepet, 2008; Takahashi et 

al., 2008; Friis et al., 2011). Fossil Lagerstätten, 

amber deposits, and insect damage found on fossil 

plants have been shown to document plant-insect 

interactions, including pollination, herbivory and 

palynivory, insect mining and galling, and insect-

plant mimicry (Wilf and Labandeira, 1999; Wilf, 

2008b; see art in Wang et al., 2012b, 2014; Bao et 

al., 2019; Correia et al., 2020; Cariglino et al., 2021; 

Tihelka et al., 2021; Xiao et al., 2021; Prevec et al., 2022). 
Plants are emerging from the background of 

ancient ecosystems in modern paleoart. The 

Ancient Colorado and Ancient Denvers murals 

and related museum reconstructions accurately 

reconstruct the history of the Denver Basin 

based on decades of detailed stratigraphic, 

paleontological, and paleobotanical research 

and collaboration with artists and sculptors 

(commissioned by Kirk Johnson and the Denver 

Museum of Nature and Science, and brought to 

life by artists Jan Vriesen, Donna Braginetz, and 

Gary Staab; Johnson and Raynolds, 2006; Johnson 

and Stucky, 2006). These murals reconstruct 

ancient environments from specific fossil 

localities, instead of broad summaries of entire 

time periods that tend to depict plants and animals 

in the same reconstruction that did not actually 

coexist (common in 20th-century paleoart). 

Some of the exceptional plant-centered artwork 

of Smithsonian scientific illustrator Mary Parish 

includes the floristic turnover of the Carboniferous 

Rainforest Collapse and the vegetation of the 

latest Cretaceous (Montañez, 2016; Sutton, 2019). 

The murals of Jay Matternes expertly recreated 

the ecosystems of North America throughout the 

Cenozoic, detailing the diversification of modern 

mammal lineages and the rise of grasslands 

(Carrano and Johnson, 2019). By assembling 

detailed geochemical, stratigraphic, and 

palynological data, Klages et al. (2020) together 

with artist James McKay illustrated the once-

diverse late Cretaceous polar forests of Antarctica 

(Figure 1F). Even traditional vertebrate-centered 

paleoart is often more conscious of the plant 

constituents than similar art 20 years ago (Figure 

1D). In recent documentaries, video games (e.g., 

Saurian, Urvogel Games), and comic books, the 

vegetation is carefully considered to reflect the 

fossil record of the time period and region (Ehret, 

2019; Parker, 2021; Clements et al., 2022; Wings et 

al., 2023). 
Among the resources available for plant 

paleoartists, the Extinct Plant Paleoart Database 

(Jud, 2020) collects examples of published paleoart 

in an accessible and continuously updated format. 

The database currently includes 177 references 

to plant paleoart, as well as a separate list of 

plant paleoartists. Although the issue of paywalls 

associated with scientific journals still hinders 

full accessibility to paleoartists, this represents 

an important first step to increase visibility of 

available resources. We hope that these recent 

scientific and artistic advancements encourage 

paleobotanists to continue collaborating with 

artists in their research and engagement to reduce 

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plant blindness and inspire future generations of 

paleobiologists to study extinct plants and animals. 

ACKNOWLEDGMENTS

E.J.S. and L.A.G. thank Peter Wilf for in-depth 

discussions on this topic as well as Cassandra N. 

Nuñez Sanchez, Rebecca Horwitt, Linda Musser, 

and the Pennsylvania State University Libraries. 

E.J.S. and L.A.G. are grateful for the fruitful 

discussions in the Pennsylvania State University 

Paleobiology Seminar and Paleobotany course on 

these topics. M.C. thanks Nathan Jud, Rebecca 

Dart, Ida Kalsta, Julianne Kiely, and Dolev 

Fabrikant for discussions on the topic. S.L. thanks 

Chris Manias and the Popularizing Palaeontology 

collective for an invaluable forum to discuss this 

topic. We are also grateful for thoughtful feedback 

and suggestions from two anonymous reviewers. 

We acknowledge financial support from NSF 

Grants EAR-1925755 (to E.J.S. and L.A.G.) and 

DGE-1255832 (E.J.S.). 

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