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Teaching physiology to the extreme: learning through human outliers 1 2 Madison J. Fry, Zachary A. Riley 3 4 Department of Kinesiology, School of Health and Human Sciences, Indiana University-5 Indianapolis 6 7 8 9 Corresponding Author: 10 11 Zachary A. Riley, zariley@iu.edu 12 13 14 New and Noteworthy 15 We discuss a novel way of delivering physiology education based loosely on the previous 16 discussion of superhero physiology. Rather than using examples that are not realistic, we use 17 examples of human outliers. 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 mailto:zariley@iu.edu Abstract 47 There are many ways to deliver physiology education as an interdisciplinary subject, yet 48 a lot of them can be dry or considered boring by students due to the subject matter. Several years 49 ago, the idea was introduced of teaching physiology based on superheroes. Learning physiology 50 in the context of these mythical people with superpowers is much more stimulating. However, it 51 lacks a translational component because humans are not faster than a speeding bullet or stronger 52 than a locomotive. We have taken this same basic concept and applied it to the extremes or 53 outliers of human physiology. What is unique or special about the physiology of individuals like 54 Lionel Messi? Michael Jordan? Usain Bolt? We give examples of two research reviews that we 55 examine in a graduate course on physiology and also provide a framework for how the content is 56 discussed in the classroom. 57 58 59 60 61 62 63 64 65 66 67 68 69 Introduction 70 71 There are hundreds of ways to deliver physiology-based curriculum to undergraduate and 72 graduate students. Many are based on pedagogy methods such as active learning (1, 2), 73 experiential learning (3), and teaching simulations (4), while many others are still centered 74 around traditional lecture-based delivery. While the teaching method is important, the content is 75 far more important. To that end, physiology as a subject-matter has had an identity problem for 76 many years, simply because it is an interdisciplinary science (5). Whether it is at the level of the 77 system, cell, or molecules, the analogy exists that they are all examining the same camera photo, 78 just subject to different levels of zoom. Unfortunately, many students studying physiology in the 79 current environment get bored staring at the camera photo, no matter what level of zoom, and 80 would instead rather be staring at TikTok, Instagram, Snapchat, or other social media (6, 7). 81 82 Superhero Physiology 83 As physiology educators, we can complain about the brain drain of social media, or we 84 can put forward a competitive alternative for the attention of the student. We cannot change the 85 content that makes up the interdisciplinary field of physiology, but we can change the context 86 through which it is applied. There is no better example of this than when Zehr first published his 87 book “Becoming Batman: the possibility of a superhero” (8). Despite Batman not having any 88 extraphysical abilities or “superpowers”, it is inherently more interesting to learn about Batman’s 89 potential physiology than it is to learn about the physiology of a similar-aged, sedentary, pre-90 hypertensive, pre-diabetic, moderately overweight male who sits in a cubicle all day. From the 91 perspective of a clinician, obviously it is going to be more important to understand the 92 physiology of the latter, but it is a lot easier to grasp the concepts applied to the Dark Knight. 93 Since that time, the concept of superhero physiology has become more prevalent and been used 94 in many different contexts (9-12). 95 It is interesting to think about how Captain America’s super soldier serum and exposure 96 to Vita-Rays could have, in an instant, changed his body mass, skeletal muscle properties, and 97 cardiovascular capabilities (9). Alternatively, you can use Hawkeye’s advanced eyesight to teach 98 vision (13), or Ironman to explain brain-machines interfaces (14). Each of these capture the 99 imagination and open up much more discussion than studying a powerpoint slide with the cAMP 100 signaling pathway or Krebs cycle on it. This is not to say that learning these processes are 101 unimportant, but rather that you could learn them in the context of these superpowers. What 102 changes? What doesn’t? Where are the optimal targets for manipulation? What are the rate-103 limiting steps? As already discussed by Brown et al. (9), you can have unlimited discussions, 104 case studies, and class projects centered on a superhero, a superpower, or a single physiological 105 system that would have to change in response to these extra abilities. 106 The only one significant drawback to the chimeric world of superheroes is just that it 107 does not actually exist (not to be argumentative here). Consequently, it is difficult to relate 108 Wolverine’s rapid regeneration and immunity to the physiology of a burn victim or someone 109 undergoing chemotherapy. What makes superheroes so super is that what they do is outside the 110 realm of physiological possibility. Interestingly, according to the website Ranker, which is a 111 democratized public voting website, Batman is the top ranked superhero of all time in comic 112 books (www.ranker.com). To obtain that ranking is quite a feat, particularly for someone who, as 113 mentioned before, has no actual superpowers other than wealth and potential influence. What we 114 might be able to take away from this is that people are more interested in what may be 115 considered optimal physiology, rather than superhero physiology. What is obtainable, even in the 116 world of fiction and fantasy as opposed to what is completely unobtainable. 117 118 Superheroes or Mere Mortals? 119 Many years ago, we implemented an alternative approach to teaching physiological 120 principles to graduate students based loosely on the idea of superhero physiology. Our approach 121 was to consider the limits of what is physiologically possible, or at least documented in the 122 scientific literature, and to use that as our model for teaching this interdisciplinary subject. This 123 does not include anecdotal accounts of “hysterical strength” such as lifting a car off a child. 124 Instead, we focused on accounts from society, hobbies, and sports that have resulted in 125 physiological extremes, or outliers. What has enabled Michael Phelps to achieve 23 Olympic 126 swimming medals, or Hafthor Bjornsson to deadlift 505kg off the ground? As to not focus 127 exclusively on sports, what enables individuals to live well past 100, or survive and thrive in 128 extreme conditions such as Sherpas on Mt. Everest? These scenarios, and many others, provide a 129 look into the unique and adaptive physiology that these individuals possess. Like superhero 130 physiology, they create an interest that is difficult to match with “normal” individuals. 131 Over the last 10 years, we have curated a series of scientific review articles (12, in total) 132 that fit the mold of the content we are trying to teach, while also being relevant and interesting 133 enough to leave the students wanting to learn more. For the sake of this article, we will be 134 summarizing two review articles that are read in the class. We will discuss the main thesis of 135 each article and then touch on some of the physiological processes that can be explained with 136 each. It is important to note that over the time the course has been delivered, we typically switch 137 out 2-3 articles as we find other research that may be more appropriate for teaching the concepts 138 we want to include. However, these are twothat have stayed in the rotation for several years and 139 are a good representation of what we are wanting to display. The articles are distinct and provide 140 different viewpoints of similar physiological processes. The first is the traditional view from the 141 lens of extreme athletic performance, while the second discusses physiological processes shaped 142 by environmental and ecogeographical factors. 143 144 Running Towards the [Almost] Impossible 145 One of easiest ways to explore physiological extremes is in the world of sports. Some of 146 the greatest feats we see in sports today occur in the world of distance running. In fact, in 2019 147 the world saw Eliud Kipchoge run a sub-2 hour marathon and in 2025 Faith Kipyegon attempt to 148 break the 4 min mile as a female. These two athletes are accomplishing feats once thought 149 impossible and they share one thing in common; they are both Kenyan. What makes these two 150 athletes, or certain countries dominate the game of distance running? The review written by 151 Wilber and Pitsiladis (15) points out that there are multiple physiological factors (genetic, 152 cardiovascular, metabolic, and muscle) that contribute to the success of Kenyan and Ethiopian 153 distance runners, and it provides a great basis for discussing how these factors have evolved and 154 change with time and training. 155 156 1. Genetically, the success of Kenyan and Ethiopian distance runners is thought to be due to 157 multiple advantageous genes as no single gene polymorphisms studied seem to provide 158 them with an advantage over other elite athletes or even the general population. However, 159 this gives the opportunity to discuss the distinct haplogroups that make up different 160 regions of the world and characteristics that are present in each. 161 162 2. Kenyan and Ethiopian distance runners have elite level maximal oxygen uptakes 163 (VO2max) as studies have reported VO2max levels between 70-80 mL/kg/min (16, 17) . 164 However, the paper reports that these values are not significantly different from elite 165 distance runners in other countries, despite Kenyan and Ethiopian distance runners 166 posting much faster distance racing times. Kenyan and Ethiopian runners also have, on 167 average, higher hemoglobin concentrations and hematocrit levels (16.4 g/dL and 49%, 168 respectively). These higher levels allow for better oxygen utilization and oxygen carrying 169 capacity. These two mechanisms can be discussed in detail, as there is a long-standing 170 debate in the relationship between VO2max levels, hemoglobin concentrations, and other 171 topics such as hemodilution or anemia. 172 173 3. Kenyan runners have an ideal ectomorphic somatotype (slim, longer, and lighter legs) 174 which allows them to be more biomechanically and metabolically economical. 175 Muscularly, they also have a higher distribution of type-I fibers (at least in the 176 gastrocnemius and vastus lateralis) contributing to greater muscular endurance. These 177 physiological characteristics can allow us to discuss the integration of cardiovascular and 178 hematological advantages and how they transfer to movement of the human body. 179 180 What truly makes this population different is their living and training environment. Kenyan 181 and Ethiopian runners grow up at altitude in an economic and geographical environment where 182 their standard travel method is often running. This has led to a myriad of physiological 183 differences in competitive Kenyan and Ethiopian runners most people cannot replicate. For the 184 sake of comparison, the average male has a VO2max of ~33 mL/kg/min (18), an endomorphic 185 somatotype (19) leaving less room to be biomechanically and metabolically efficient, and have 186 approximately 41% type-1 fibers in their vastus lateralis (20). The great feats of Kenyan and 187 Ethiopian distance runners are nearly impossible for the average human to achieve and provide a 188 great model to understand outliers in human physiology. 189 190 Allen’s Rule: Trying to Stay Cool 191 192 Different from exploring physiology through an internal genetic and biologic lens, 193 physiology can also be looked at through an external environmental lens. Joel Allen took this 194 approach when he came up with his own rule that the environment and ecogeographic location in 195 which we live (ambient temperature and latitude to the equator) plays a role in extremity length 196 relative to the trunk (21). The review article by Pomeroy et al. (22), continues to explore the idea 197 of Allen’s rule, while also exploring the influence of population history and environmental 198 stressors (pathogen load and resource availability) on human stature and provides a great basis 199 for discussion on if this rule has held up over time and how it relates to thermoregulation, 200 metabolic efficiency, and developmental plasticity. 201 202 1. Being able to maintain a constant internal temperature in any environment is an important 203 negative feedback process in maintaining homeostasis. Due to this, thermal adaptations 204 are thought to play a significant role in human stature and morphology. Longer limbs 205 provide a greater surface area for evaporation, allowing for better maintenance of 206 homeostasis in hot environments. Conversely, shorter limbs and greater body mass 207 provide a decreased surface area allowing for the conservation of heat in cold 208 environments. This thermoregulatory framework allows us to discuss why some 209 morphologies are better suited for certain climates and how the body can adapt, even with 210 a less-than-ideal morphology, in a different environment. 211 212 2. As previously stated above, thermoregulation is important for maintaining homeostasis, 213 but that does come at a metabolic cost. Having short limbs allows individuals to be more 214 metabolically efficient at least in terms of thermoregulation compared to those with long 215 limbs resulting in greater heat dissipation (23). This opens discussion on how important 216 human stature and body composition is on the metabolic efficiency of sport performance 217 or general living in different environments. 218 219 3. Environmental temperature and stressors (i.e. nutrition and pathogen load) have been 220 shown to affect development plasticity. Warmer environments have been shown to 221 increase bone growth comparative to cold (24, 25). Nutritional inadequacy and 222 environments with higher pathogen loads have been associated with plastic responses in 223 human stature and relative lower limb length showing a negative relationship. This 224 information provides the opportunity to discuss how the environment and genetics can 225 affect the phenotypes that are expressed during development and what happens if certain 226 phenotypes are expressed differently due to stressors. 227 228 This provides further understanding and discussion about how the world and climate we 229 live in affect human physiology. A great example, relevant to this paper and also the paper from 230 Pomeroy et al. (22), are men from the Dinka tribe in South Sudan. These individuals, including 231 former NBA basketball player Manute Bol, are some of the tallest in the world with the longest 232 limbs, allowing for greater heat dissipation. There are many different factors that contribute to 233 the unique phenotype of the Dinka tribe, and from a teaching perspective the discussion can 234 include their genetic cluster in Africa, general nutrition and nutrition availability, as well as the 235 effects of poverty, disease, and the introduction of modern healthcare. 236 237 Course Delivery 238 As seen above, each of these articles offer a unique perspective on physiological 239 processes. The goal remains consistent: toentice or enthrall the students in a manner similar to 240 studying superhero physiology, but to present it in a way that is realistic and physiologically 241 possible. Now that the content is clarified, we need to consider how it is conveyed in the 242 classroom. Each classroom session is broken up into two main activities: 1.) Student group 243 presentations over the assigned review paper, and 2.) roundtable discussion over the assigned 244 review paper and other tangential topics. We will discuss each of these in more detail below. 245 Each review paper is assigned to a group of 3 students, with the expectation of them 246 delivering a 45-minute powerpoint presentation over it. The presentation instructions provided to 247 the students are shown in Figure 1. The rationale behind having the students present the material 248 is two-fold: 1.) it is a low-risk environment to develop research and teaching presentation skills 249 to help mitigate public speaking fears that students may have (26), and 2.) it increases the 250 students’ exposure to academic and research language and improves scientific literacy through 251 practicing science-process skills (27, 28). The latter could be partially addressed by just having 252 the students read the research and write up a summary or report, however, they would miss out 253 on the benefits of retrieval practice, which is known to aid in material retention (29). As for the 254 former, it is well accepted that you only get better at presenting (and reducing public speaking 255 anxiety) through practice and repeated exposure (30). 256 Insert Figure 1 About Here 257 258 Leading into the week’s presentation, the students that are not presenting are expected to 259 read the paper and come to class with three prepared questions for the roundtable discussion. The 260 only rule given on the questions is that they cannot be answered with one word or number (e.g. 261 ‘yes’ or ‘no’) and the students hand in their questions at the end of class to count for an 262 attendance grade. 263 264 Examples of student questions from Wilber and Pitsiladis (15): 265 1. With the extensive walking and running, for example to and from school, do Kenyan 266 distance runners have lower perceived effort at a given level of intensity? 267 2. Can the success of East African runners be replicated in other high-altitude regions? What 268 if the altitudes are even more extreme, e.g. Himalayas? 269 3. How do mitochondrial density and oxidative enzyme activity contribute to the superior 270 endurance capabilities observed in these athletes? 271 4. How does a lifetime of running barefoot, or with minimal footwear, impact running 272 economy? 273 5. How do hormonal profiles (e.g. cortisol, testosterone) differ in these runners due to 274 chronic training and continuous altitude exposure? 275 276 Following the presentation, the students are organized in a circle for a roundtable discussion 277 based on the Socratic method of teaching. The application of the Socratic method in our 278 roundtable discussion relies on a few key points illustrated by Delic and Bećirović (31): 279 1. We as teachers are not teaching, but rather facilitating and guiding. 280 2. We avoid “rights” and “wrongs” and instead force them into using reasoning based on 281 prior knowledge and experience. 282 3. Encourage students to follow different paths of logic to strengthen their own 283 understanding. 284 These practices are emphasized to turn the discussion over to the presenters with the most 285 intimate knowledge, and their classmates that have prepared questions, in advance. There are 286 some additional tips that we have learned in the past decade that can be used for engaging the 287 class in discussion. If a student is not comfortable sharing a question that they wrote, instead ask 288 them to share their opinion on a question that is already being discussed. Encourage students to 289 have their electronic devices out and ready. There are many times when scenarios are brought up 290 that additional information can be gleaned from a simple google search and then shared with the 291 class. Host the class in a room that has a whiteboard, as some students feel more comfortable 292 drawing or writing out their understanding of a system or mechanism. Finally, do not be afraid to 293 role play to have the students think from a different point-of-view, such as an athlete or coach. 294 295 This leads to the obvious question of “what do the presentations and roundtable 296 discussions based on the Socratic method have to do with teaching the physiology of extremes?”. 297 If you have ever tried to turn a class discussion over to students when the material is dry and 298 boring, you quickly realize that even the brightest extroverts suddenly become quiet. Instead, 299 using review articles that discuss physiological extremes, such as those presented above, creates 300 interest among the students and leads to a number of “what if?” questions. The “what if?” 301 questions from students are perfect for the roundtable environment because seemingly everyone 302 will have an opinion or different take. As a teacher, all we then need to do is follow the three 303 principles listed above for implementing the Socratic method. This is not failproof, as no 304 teaching strategy is, but it seems to be appropriate for the content and topics that we cover. 305 We have not collected formal quantitative data on student-learning outcomes, but we do 306 have student evaluation scores from the 5 times this class was offered over the last ten years. 307 That data is presented in Table 1 next to the department averages for those same semesters. There 308 were 4.04%-7.65% increases in the different student evaluation categories over the departmental 309 averages for this class. We have also documented feedback from the written comments in the 310 student evaluations. Below are a couple of comments received over the last 10 years: 311 “This course was very rewarding and allowed for the students to have round table discussions on 312 topics related to the various fields we may enter as well as allowed us to think critically and 313 discuss in a safe space.” 314 315 “I really enjoyed the layout of this course. Learning in a discussion-based setting really helped 316 me think more critically on the topics presented. It got me excited about research and academia 317 in general.” 318 319 There have also been comments that were useful in helping us make improvements in the 320 class. 321 322 “If possible, let the students pick a topic and paper for their 3 rd and final presentation” 323 324 “Pacing of class could start slower; adding in the instructor in the first one or two presentations 325 could allow students to be better prepared before each class for discussion and be able to 326 dedicate more time to being able to produce more in–depth and better presentations.” 327 328 Both comments will be addressed the next time the course is delivered. First, we will 329 allow students to pick their research topic for their final presentation. We will likely need to vet 330 the studies and topics to make sure they fit the class objectives, so we will ask them to bring 331 three different papers in for one to be approved. Second, the instructor will deliver the very first 332 presentation and lead the first discussion so that the students have an example to use when 333 preparing their own. 334 Insert Table 1 About Here 335 336 Conclusion 337 There is a reason why the Guinness Book of World Records has sold over 155 million 338 books globally (guinnessworldrecords.com). People are always interested in the extreme, record-339 breaking, or absurd. As many authors have previously stated, this is what has made superhero 340 physiology such an interestingconcept. We have attempted to take a similar approach to teaching 341 physiological processes, but have stayed grounded in what is possible, attainable, and 342 observable. 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European 423 Researcher 111: 511-517, 2016. 424 425 426 427 428 429 430 431 432 433 434 Table Legends 435 Table 1. The average course evaluations for the Department and for this course for the 5 times it 436 was offered (every two years) in the past 10 years. The general themes that were evaluated are 437 listed in the left column. “5” is strongly agree and “1” is strongly disagree. 438 439 Table 1. 440 Dept Avg (/5) Phys Avg (/5) % Increase Course 4.54 4.72 4.04% Instruction 4.48 4.75 5.95% Learning 4.52 4.81 6.42% Assignments 4.51 4.72 4.73% Communication 4.49 4.83 7.65% 441 442 443 Figure Legend 444 Figure 1. Instructions provided to the students for the presentation requirement. 445 446 447 448 449 450 451 452 Article File Figure 1