Keywords

1 Introduction

To explain why I work on honey bees, I first want to tell you a little bit about my journey as a biologist. Biology is often portrayed as a continuum made up of different levels of organization, from cells to tissues, to organs and organ systems, to individual organisms, and to populations, communities, and ecosystems. Wikipedia defines cell biology as the “branch of biology that studies the different structures and functions of the cell and focuses mainly on the idea of the cell as the basic unit of life” (Wikipedia contributors, 2018). As an undergraduate majoring in Biology at Williams College, I became fascinated with the cellular and molecular aspects of biology. As part of a continuum, the concept of cell biology in isolation is relatively abstract without attempts to integrate it with broader biology, chemistry, and physics. Especially when thinking about multicellular organisms, cell biology is most relevant when it can be applied to understanding higher levels of organization. The cell biology definition in Wikipedia continues on to suggest that “knowing the components of cells and how cells work is fundamental to all biological sciences; it is also essential for research in biomedical fields such as cancer, and other diseases” (Wikipedia contributors, 2018). True to form, I was thinking about how biology at this level contributed to disease in humans. After graduation, I studied mammalian blood and immune development and function. I was involved in looking at the cell biology of how we make our red blood cells, our white blood cells, and our platelets, often in the context of blood disorders and malignancies. Primarily focusing on mouse models and cell lines, I examined the role of signaling pathways and transcriptional regulators affecting these processes in both my Ph.D. and post-doctoral work. The relevance of my studies to disease in humans was always in the backdrop.

During my postdoctoral training, I had the opportunity to teach a course on the American food system with a longtime friend. This served as the beginning of a continuing interest in the systems, and especially the organisms, involved in making our food. As I looked towards the future, I hoped to use the techniques and ways of thinking I had learned in my graduate and postdoctoral training with a new model organism that might bring together these interests. With bees, I felt perhaps I had found the perfect model. Bees are fascinating and they are currently experiencing a crisis, but we do not know much about them at the cellular and molecular level. Luckily, however, their genome is sequenced and we can use knowledge gleaned from other model insects to inform our questions. Perhaps most importantly, a cell biological understanding of biology is highly transferable between animal species, so I began to use my skills in this new area.

While honey is one of the most commonly thought of benefits supplied by these insects, the most important benefit is the pollination services they offer (reviewed in Snow & Rivera Maulucci, 2017). The Western Honey Bee, Apis mellifera, provides pollination services of critical importance to humans in both agricultural and ecological settings (Goulson et al., 2015). When bees fly from flower to flower foraging for nutrients, they transfer pollen between flowers of the same species allowing for fertilization of the ovule and completion of the plant’s reproductive cycle (see Fig. 8.1).

Fig. 8.1
A photograph depicts a honey bee pollinating a cherry blossom.

Honey bee pollinating a cherry tree on Riverside

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The co-evolution of pollinators and plants over millions of years has made bees and other animals a critical part of many ecosystems (Potts et al., 2016). In addition to their importance in natural ecosystems, this pollination service is absolutely required for the growth of many agricultural crops important to humans.

Plants like almonds absolutely require insect pollination. As our agricultural systems have become more homogeneous, and monocultures have become the norm, honey bees have become the only pollinator with the ability to effectively pollinate many of the crops in our modern agricultural system. Some examples include apples, cranberries, melons, and broccoli with some crops like almonds, blueberries, and cherries being greater than 90% dependent on honey bee pollination. Honey bees are easy to move, provide a high number of individuals, and they are generalists in the plants they will pollinate. To satisfy the pollination needs of an industrialized agriculture system, “industrial beekeeping” practices are now employed. According to a 2015 survey, over 85% of the country’s 2.6 million colonies were in commercial migratory beekeeping operations with an average of >4000 hives (Lee et al., 2015). Recently, meeting the pollination demand has become even more challenging as honey bee colonies have suffered from increased mortality, most likely due to a complex set of interacting stresses (Goulson et al., 2015). Key stresses thought to be involved include nutritional stress due to loss of appropriate forage, chemical poisoning from pesticides, changes to normal living conditions brought about through large-scale beekeeping practices, and infection by arthropod parasites and pathogenic microbes (see Fig. 8.2).

Fig. 8.2
A photograph represents a honey bee, where key stressors that impact bees are labeled management stress, chemical stress, nutritional stress, infection stress, and climate stress.

Key stressors impacting honey bee health

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The challenges facing pollinators mirror those facing diverse organisms in both natural and managed environments. These issues will have far ranging impacts on the health of individuals and communities and the negative effects will be felt disproportionately by marginalized and disadvantaged groups. For example, decreased abundance and diversity of animal pollinators like bees (Ollerton, 2017) is predicted to have potentially dire consequences for food security (Smith et al., 2015) that will be felt especially acutely in areas that are ‘food deserts’ where little food stuffs are produced, such as urban centers (Lawson, 2016).

As no single cause for the recent increase in honey bee disease is evident, there is enhanced focus on the impact of interactions between various stressors. In seeking to understand how stresses might synergize to impact honey bee health, we have undertaken efforts to more completely define common cellular processes and cell stress pathways that are impacted by multiple stressors. My lab studies cell stress and infectious disease in honey bees. Undergraduates are incorporated into every aspect of this research throughout the year, with particular emphasis on designing and performing experiments as well as analyzing data. Figure 8.3 shows undergraduate students working with the bees.

Fig. 8.3
A photograph depicts five people dressed in snow lab coats and masks, where a man holds the honey bees in the bee box.

Snow lab members installing a ‘package’ of bees

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Together, we are interested in two big questions. First, how do bees respond to stress at the cellular level? Second, how does one of their common parasites, Nosema ceranae, respond to stress at the cellular level? We hope to use the answers to these two questions to improve honey bee health.

We are specifically focused on the process of proteostasis, which refers to the homeostasis of protein synthesis, folding, function, and degradation (reviewed in Taylor et al., 2014). In multiple organisms, a number of normal and pathologic conditions shown in Fig. 8.2 can lead to disruption of proteostasis. Environmental factors, as shown in Fig. 8.4. can modify proteostasis by acting through these cellular and organismal pathways, including microbial infection, aging, and nutrition, all of which are implicated in honey bee health.

Fig. 8.4
A circular model represents proteome function, which includes proteostasis modifiers, proteostasis network signalling pathways, and proteostasis pathways.

Proteostasis is regulated at many levels of biology. (Adapted from Powers & Balch, 2013)

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Work in other organisms has defined a number of cellular pathways that are involved in recovering from proteotoxic stress, including the Unfolded Protein Response (UPR) centered in the endoplasmic reticulum (Walter & Ron, 2011) and the Heat Shock Response (HSR) centered in the cytoplasm (Vabulas et al., 2010). In our previous work, a number of very talented undergraduates have worked to characterize these two proteostasis pathways (Johnston et al., 2016; McKinstry et al., 2017; Shih et al., 2020, 2021; Adames et al., 2020; Bach et al., 2021; Flores et al., 2021). These studies are ongoing with new cohorts of students helping to define unique aspects of honey bee cell stress pathways and the stress pathways of their pathogen, N. ceranae (McNamara-Bordewick et al., 2019).

In the context of a research laboratory, it has been relatively easy to discuss the plight of the honey bee with students and to draw broad links between our work on cell biology and larger issues of sustainability. This has been, in part, because research students have direct interaction with bees; helping to tend the hives, assisting in extracting honey, collecting and caring for bees in caged experiments, and dissecting specimens to obtain samples (see Fig. 8.3). In the next section, I will talk about my efforts to bring sustainability in thought and action into my classroom, with specific focus on my course, Laboratory in Cell Biology.

2 Sustainability in the Classroom

As a faculty member at Barnard, I have been fortunate to teach exceptional students. I have drawn on my experiences as an undergraduate at Williams College, where I was nurtured in both intellectual creativity and curiosity as well as challenged to develop sharp critical-thinking skills. I was hired as a cell biologist and I am primarily responsible for teaching three classes that deal with this level of biological organization, Laboratory in Cell Biology, Cell Biology, and Introduction to Cell and Molecular Biology. I come from a biomedical background and am still quite interested in questions of human health, but as described above, my big picture view has shifted to focus on the environment and sustainability, especially as they connect with agriculture and our food production system.

My first thoughts about making a more intentional effort to incorporate sustainability into my classrooms came from a collaboration with María Rivera Maulucci in the Education Department here at Barnard. We co-wrote an article about using honey bees in education (Snow & Rivera Maulucci, 2017) and it turned out to be a fun, educational, and inspiring experience that allowed for considerable self-reflection and a renewed desire to merge sustainability and teaching more purposefully. This experience synergized with my concurrent involvement with the Sustainable Practices Committee at Barnard. One of the activities of this committee was to hold a workshop series during the 2017–2018 academic year, in which the college community gathered to work towards the creation of a shared climate action vision statement. The workshops focused on a variety of areas, including Consumption and Waste, Energy, Local Environment, Curricula and Research, and Campus Culture. One of the goals of these workshops was to encourage faculty members to envision ways to include sustainability into our broader curriculum.

During the campus conversation on Curriculum and Research, María introduced us to Banks’ ideas about incorporating multicultural concepts into a curriculum (Banks & Banks, 2016). She suggested that Banks’ framework could be applied to introduce sustainability into a classroom (Rivera Maulucci, Pathways, this volume). The strategies proposed by Banks, and adapted by María, allow for different amounts of integration of this new perspective, in this case sustainability, with the ‘core’ material of the class. The ‘levels of integration’ proposed by Banks include: The Contributions Approach (Level 1), The Additive Approach (Level 2), The Transformation Approach (Level 3), and The Social Action Approach (Level 4). According to Banks, Level 1 would see inclusion of a small number of discrete exposures to the new perspective without substantial connection to the existing course material. Level 2 would incorporate the concepts associated with the new perspective into the whole course without a substantial change to the course structure. Level 3 involves a major restructuring of the curriculum with the intent to fully integrate new material. Level 4 encourages students to apply the new perspective to real-world issues and pursue activism to support social change shaped and supported by what they have learned. Thus, the intensity of sustainability inclusion is dependent on the flexibility of the original material, the teacher, and the students involved in the class.

The collaboration with María and the work as part of the Sustainable Practices Committee helped me to imagine incorporating sustainability more fully into one class I teach, Laboratory in Cell Biology. In the case of this class, the broader context of the material is quite flexible, so I could employ the more intensive Transformation Approach (Level 3) and restructure the curriculum to fully integrate sustainability. In addition, although the students are Biology and Neuroscience majors primarily interested in health-related fields, they are some of the best and the brightest STEM students at Barnard, and I knew they would be ready to meet any challenge. To describe the proposed changes, I first want to start by describing the class and giving some context on how Barnard’s Biology Department envisions laboratory classes.

Research on science pedagogy has revealed that involvement with the theory, practice, and culture of science as undergraduate researchers increases long-term persistence in STEM (Linn et al., 2015). Providing such experiences is a major goal of a research program at a small liberal arts college. In my research lab, student researchers play a significant role in working to elucidate cellular stress responses of honey bees. However, as the number of students that can be included in a research program at a small college is necessarily limited, another approach to exposing a broader set of students to the ‘authentic’ theory, practice, and culture of science is through the use of upper-level labs. Authentic learning is an instructional approach that allows students to apply their knowledge to real-world problems while modeling the theory, practice, and culture of professionals in the field. Important aspects of authentic learning include open-ended inquiry, learning in groups, and the use of independent project work (Rule, 2006).

In our department, there has been a major push to implement these authentic learning principles in lab classes, mirroring a trend that has been gaining popularity in science education for some time nationally. One model for achieving these goals in known as Course-Based Undergraduate Research Experiences (CURE) (Auchincloss et al., 2014) and is defined as a course in which “students work on a research project with an unknown answer that is broadly relevant to people outside of the course” (Cooper et al., 2017). While not a CURE class in the strictest sense, the lab class I teach, Laboratory in Cell Biology, incorporates many of its principles, pursuing cell biology through weekly lab experiments that allow about 16 upper-level students to gain experience in the theoretical and practical aspects of how cell biologists think about and perform experiments, collect data, and present scientific findings. The incorporation of original research questions interconnected with those pursued in my research lab represents a key aspect of this class that increases the engagement of students by incorporating “authenticity” into laboratory class. The inclusion of an independent project for the last third of the class is especially valuable in helping meet the goals of authentic learning. In addition to providing an authentic research experience, classes based on the CURE model have been shown to help increase the participation and retention of underrepresented groups in STEM education (Rodenbusch et al., 2016). Thus, CURE-type classes can be a major force for increasing inclusion in the science education and hopefully in the scientific endeavor more broadly.

When I first developed the class, I decided to focus on a small part of cell biology, specifically choosing an area that was directly related to my research; proteostasis and the cellular responses to disruption of this process. As mentioned above, proteostasis is maintained by the responses of the proteostatic network within individual cells. These pathways sense proteostatic stress through the detection of a build-up of unfolded proteins and trigger a suite of responses designed to return the cell to homeostasis. Three highly conserved pathways are responsible for sensing proteotoxic stress in the endoplasmic reticulum and trigger the coordinated response known as the UPR (Fig. 8.5). These pathways are characterized by unique transmembrane receptors and signal transduction machinery which activate UPR target genes to return the cell to homeostasis (see Fig. 8.5).

Fig. 8.5
A model represents unfolded protein pathways, which include I R E 1, P E R K, A T F 6, Active X B P 1 S, A T F 4, A T F 6, and Active X B P 1 S, A T F 4, A T F 6.

The unfolded protein response pathways

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For the class, we use a fruit fly (Drosophila melanogaster) cell line (Schneider, 1972), because they are easy to grow and many protocols exist for experimenting with them. But as a fruit fly-derived cell line, they are far removed from bees and pollinators. For framing and readings, I have focused on the cell biology and human diseases associated with the dysfunction of the processes we study, such as Alzheimer’s (Hetz et al., 2020). Because of the flexibility of the cell line model, we can easily look at the process of proteostasis and the response to it by measuring various steps in the process including signaling, changes in gene expression, and cellular outcomes (Samali et al., 2010). Based on my assessments and student evaluations, this class has been quite successful in its aims of introducing the theoretical and practical aspects of how cell biologists consider their field while increasing student enthusiasm and engagement. However, most students consider the broader biological context of the class to be about human disease, with approximately 40% of biology majors entering MD programs (for years 2010–2015). As biology is a continuum made up of different levels of organization and cell biology is highly transferable between metazoan species, it should be relatively straightforward to change the context to one involving themes of sustainability.

I devised five strategies to help link cell biology to problems impacting honey bees (see Table 8.1). First, I framed the class in terms of pollinators and bee health. Instead of beginning with human diseases, I started with pollination, honey bee biology, and honey bee health. Readings now reflect this change in framing (e.g., inclusion of Potts et al., 2016). I still make connections with medicine, but by altering the structure to move medicine to later dates in the course, I can shift the emphasis to sustainability. Second, I established this new emphasis by bringing live honey bees into class on the first day (with trips to see the apiary on the roof when weather permits). Third, I incorporated original research from my lab into the readings. Previous work examining proteostasis pathways in honey bees have resulted in publications with student coauthors that can be used in the class, including those cited above (Johnston et al., 2016) and (Adames et al., 2020). Fourth, we started with cell biology experiments using honey bees before focusing on cells. By getting to know the organisms before thinking about cells, I hoped to move through the levels of biological organization from organisms to tissues to ultimately zero-in on cells and the molecular component involved in their function. Finally, I encouraged students to explore links between cellular biological organization and sustainability problems that affect pollinators in their final projects.

Table 8.1 Planned steps to connect cell biology to pollinators
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After implementing the majority of these changes for one semester in the fall of 2018, I found the project was overall a success with some key additions to be undertaken in subsequent years. Anecdotally, I found that students were excited by the inclusion of sustainability concepts in the course and that students felt like the structural and content changes introduced them to new ideas and connections between cell biology and sustainability that they would not otherwise have been exposed to. However, I felt that there were two aspects that could be strengthened in future iterations of the course. First, there were a number of missed opportunities to solidify the link between cell biology and sustainability as the class drew to a close. I find that the themes and information in a class often fall into place for students during the last few weeks of the course when they look back and synthesize information from the whole course. This is especially true when there is a cumulative assessment, which we have in this class in the form of both a final lab report and a lab final exam. Thus, I planned to spend additional time emphasizing the themes of sustainability covered throughout the course during the retrospective/synthesis period leading up to these assignments. In addition, I aimed to explicitly include these sustainability themes into the two final assignments to encourage student engagement through the end of the term. Second, any discussion of pollinators can be directly connected to concepts of environmental justice (Mohai et al., 2009). Making those associations in the class might represent an ideal opportunity to connect science and broader societal issues in the biology curriculum. Third, I did not have a method in place for gauging the success of the project in a more concrete and quantitative manner. In order to assess the success of using the Banks framework to bring sustainability in thought and action into the Laboratory in Cell Biology class, I hoped to solicit feedback in a more structured manner.

I have now taught the class for the fourth time. A survey administered to the most recent iteration (taught fall, 2021) class assessing the effectiveness of the five strategies named above was encouraging. When asked to rate the effectiveness of the strategies, students (n = 17, Mean ± SD) gave the following scores; framing (4.58 ± 1), interacting with bees (4.71 ± 0.99), incorporating original Snow lab research (4.41 ± 1), starting with experiments on bee tissues before cells (4.65 ± 0.79), and referencing the broader context in the final projects (4.47 ± 1.07) (where 5 = very effective, 4 = somewhat effective, 3 = neutral, 2 = somewhat ineffective, and 1 = very ineffective). Feedback in comment form was quite positive (e.g. “Thank you for a wonderful class.”), but also provided some constructive suggestions for future iterations. The recommendations fell into two broad categories. First, a number of comments suggested spending more time discussing specific threats to pollinators (e.g. pesticides and infection) and bridging these insults to the UPR more concretely. A specific example was a request to emphasize Snow lab research more in recitation. Second, there were a number of calls to include even more bee species and more interactions with bees during the class the first day. For example, one student suggested using visits to the apiary and college green roofs to discuss bees, pollinators, and sustainability in a NYC-specific manner. These points represent positive and productive feedback that I hope to incorporate over the next few iterations of the class. One future goal commensurate with these student reactions is described in the next section.

3 Beyond Honey Bees

According to E. O. Wilson’s Biophilia hypothesis, humans are inherently interested in learning about other life-forms (Wilson, 1984). Many scholars and activists alike have worried about the loss of this human attribute as we have increasingly less interaction with nature in the modern world (Pyle, 1993), leading to diminishing interest in conservation (Soga & Gaston, 2016). In a 2002 article, authors described the ability of schoolchildren to name approximately 50% more Pokémon characters than real species (Balmford et al., 2002). In agreement with this finding, most students I talk to think there are only two types of bees, “honey” and “bumble.” Yet in fact there are ~20,000 other bee species on the planet (Michener, 2000). However, the vast majority of attention and resources are spent focused on the honey bee and to a lesser extent on other managed bee species (Geldmann & González-Varo, 2018). Of the myriad bee species, ~50 species are managed and only 12 are commonly used for pollination services (Potts et al., 2016). Agriculturally-important managed bees include select species of the following groups; bumble bees (genus Bombus), alkali bees (Nomia melanderi), alfalfa leaf-cutter bees (Megachile rotundata), and mason bees (genus Osmia) (Klein et al., 2018).

In addition to bees, many other animals are involved in pollination (Ollerton, 2017). A long-held belief has been that diversity in pollinators is of significant value, leading to higher levels of pollination even in the presence of managed pollinators (Kremen, 2018) and recent studies support this view (Garibaldi et al., 2013, 2016; Winfree et al., 2018). Reductions in all animal pollinators, including wild bees, have been documented in recent years (Potts et al., 2016). One study found a 50% reduction in the bee species visiting plants within a specific study region over the last century (Burkle et al., 2013). While difficult to quantify the effects, such pollinator losses could have significant impacts on human health (Smith et al., 2015). It is likely that these losses are largely due to human impacts (Winfree et al., 2011). This troubling trend of diversity loss as a direct result of our activities extends beyond pollinators to other invertebrates. For example, a 2017 study showed a 75% decrease in flying insect biomass over the last three-decade period in Germany (Hallmann et al., 2017). As a person interested in sustainability and conservation, I found these numbers disturbing, and I aspired to stretch my goals for the Laboratory in Cell Biology even further. Could I use the class to increase student’s knowledge of other bee pollinators and invertebrates more broadly? Could I use the class to think about conservation beyond species directly relevant to the well-being of human society?

Charismatic megafauna is a term used to describe widely popular animal species, which are often used by conservation advocates to achieve environmental stewardship goals (Ducarme et al., 2013). There are pros and cons to using this model to involve the wider public in conservation. However, there is some evidence that people’s natural interest in honey bees (reviewed in Snow & Rivera Maulucci, 2017) can be used to increase exposure to other wild species. Some argue that honey bee conservation on its own will not help wild bees and other pollinators (Geldmann & González-Varo, 2018) and some evidence even suggests that where introduced, managed bee colonies can have negative impacts on native bees (reviewed in Mallinger et al., 2017). In addition, it is clear that conservation of pollinators will have to be driven by forces beyond the crop pollination services these species provide (Kleijn et al., 2015). But if honey bee awareness could be the gateway to broader awareness of and interest in wild bees, other insect pollinators, and invertebrates more broadly, it could have significant beneficial consequences for the conservation of these species. For example, in March of 2017, the rusty-patched bumblebee became the first bumblebee, and only the eighth bee, to be protected under the Endangered Species Act. It is highly probable that increased awareness of the challenges facing honey bees and the importance of pollination may have contributed to this somewhat remarkable event.

To incorporate ideas of conservation beyond honey bees into this class, we are now revisiting the strategies from Table 8.1. First, when framing the class I now expand the topics to include wild bees and the challenges facing them by including additional reading about them (e.g., Klein et al., 2018). Second, when establishing the emphasis of the class we hope to include a second ‘field’ trip to the green roof located on the Diana Center here at Barnard. Wild bees are abundant in New York City (Matteson et al., 2008) and they are likely to be on the Barnard Campus on the first day of class in September. So, our first class might include a ‘bee hunt’ (Mueller & Pickering, 2010) in addition to an apiary tour. To make this exercise more meaningful, I plan to have students photo-document any pollinators they see for subsequent identification. Documentation of various organisms including pollinators has been part of multiple “citizen science” initiatives in recent years. One of the most successful has been the “bioblitz”, which involves students and volunteer experts attempting to document and identify all the species in a given area over a specific time period (Ballard et al., 2017; also see O’Donnell & Brundage, this volume). While a true bioblitz is far beyond the scope of this class, documenting pollinating insects on our green roof over a 2-h period would work well. While data from citizen science initiatives have been shown to be somewhat incomplete (Kremen et al., 2011), the impact of such programs on understanding and appreciation of species diversity and conservation are apparent (Mitchell et al., 2017). Because we do not have experts in entomology available to help with species identification in this class, we can employ a widely used crowdsourcing site, iNaturalist (www.inaturalist.org, McKinley et al., 2017) for this task. iNaturalist is described as a “crowdsourced species identification system and an organism occurrence recording tool” and allows for additional time and map data to be recorded with observations. As proof of principle, I used the site to identify the picture of the bee in Fig. 8.6, taken on the Barnard campus in late July of 2017.

Fig. 8.6
A photograph represents a bumblebee sits on a leaf.

Brown-belted bumblebee, Bombus griseocollis on Barnard’s campus. (Photo credit: Rachel Snow)

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The bee was identified by a number of experts as a brown-belted bumblebee, Bombus griseocollis, which is native to much of the United States except for the Southwest. It is my hope that even this limited exposure to wild bees and the unique challenges faced by them will shift the attitudes of the students in my class towards appreciation and conservation.

4 Full Circle from Teaching to Research

One of the many delights of being an educator is learning from your students or your own teaching. Thinking about how to incorporate sustainability into my Laboratory in Cell Biology class forced me to consider pollinators other than honey bees more than I had ever done in the past. Like with honey bee losses, native bee decreases are likely caused by a complex set of interacting stresses (Goulson et al., 2015). Yet, from a research perspective, especially at the cell and molecular level, we have, by orders of magnitude, more information about honey bees than we do about a given wild bee species (Trapp et al., 2017). One metric of the quantity of research on an organism or group of organisms is the number of publications about them. As an example, looking at specific bees, as of 2021 there are 4 scholarly publications in Web of Science about the bee species Megachile centuncularis, which is native to New York City. There are 795 publications on all bees of the genus Megachile. By contrast there are 4997 publications about the European honey bee Apis mellifera and 9521 publications on bees from the genus Apis for the last 5 years alone!

Another indicator of the degree to which an organism is being studied at the molecular and cellular level is whether its genome has been sequenced. Of the 20,000 bee species, we have genomes sequenced for only 15. I have in no way given up my interest in or commitment to honey bees. However, I hope to think about these other bee groups and be more inclusive when framing my own research questions and drawing conclusions. I have now expanded the organisms I study beyond honey bees to some wild bee species. As mentioned above, solitary bees of the genus Megachile have been studied (Pitts-Singer & Cane, 2011; James & Pitts-Singer, 2013) and tools have been developed for studying them (Fischman et al., 2017; Trapp et al., 2017). There is even some preliminary research looking at thermal stress and the immune response in these bees (Xu & James, 2012), which overlaps with the kind of cell stress pathways we typically explore in honey bees (McKinstry et al., 2017). We have now begun working with M. rotundata to better understand how what we learn in honey bees might apply to other bee species.

5 Conclusions and Reflections

Honey bees are critical pollinators for agriculture and are facing health problems of complex origin. From a research perspective, understanding them at the cellular level will be important in helping to improve their well-being and survival, with critical benefits to human nutrition. As a teacher, I am responsible for many courses focused on biology at the cell and molecular level. Biology is a continuum of different levels of organization, all of which are important and critically intertwined. Because of this, cell biology as a discipline can be framed in a variety of contexts. Although the default context is typically medicine, connecting the theory and practice of cell biology with the broader environmental issues of honey bee disease and the growing pollinator crisis is promising. Furthermore, I believe I can use the widespread appeal of honey bees to encourage students’ interest in more bee species, other pollinators, and invertebrates in general with positive impacts on conservation and sustainability. Student feedback revealed that the sustainability aspects of the class were very well received, but also provided some constructive suggestions for changes that will be the basis of future improvement to the class.