Who needs “Ice Road Truckers” when you have the “John’s Freezer” team on the road with fossils, amphibians, felids and 3D phenotype fun? No one, that’s who. We’re rocking the Cheltenham Science Festival for our first time (as a group), and pulling out all the stops by presenting two events! Here’s the skinny on them, with updates as the week proceeds.

Stomach-Churning Rating: 2/10 for now (just bones), but it could change once the cheetah dissection is under way… 8/10 bloody cheetah bits but only at the end (updated)

Right now, Lauren Sumner-Rooney (of “Anatomy To You” and other fame) is on-site with a rotating team of others from our lab, in the “Free Activity Tents” area of the Imperial Gardens/Square, inside a marquee where we’ll be showing off our NERC-funded tetrapod research all week. This “First Steps” event features not only our past and present work with Jenny Clack, Stephanie Pierce, Julia Molnar and others on Ichthyostega & its “fishapod” mates, but also our “scampering salamanders” research in Spain, Germany and England. I’ve blogged a lot about all that, and won’t repeat it here, but you can see a 3D-printed Ichthyostega skeleton, view the skeleton in a virtual reality 3D environment, see related specimens and engage in kid-friendly activities, and talk to our team about this and other related research.

Ichthyostega 3D printed backbone is born!

Ichthyostega 3D printed backbone is born!

The central themes of that event are how bone structure relates to function and how we can use such information, along with experimental measurements and computer models of real salamanders, to reconstruct how extinct animals might have behaved as well as how swimming animals became walking ones. How did fins transform into limbs and what did that mean for how vertebrates made the evolutionary transition onto land? If you know my team’s work, that encapsulates our general approach to many other problems in evolutionary biomechanics (e.g. how did avian bipedalism evolve?). Added benefits are that you too can explore this theme in a hands-on way, and you can talk with us about it in person. That continues all week (i.e. until Saturday evening); I’ll be around from Thursday afternoon onwards, too. Kids of all ages are welcome!

Ichthyostega 3D print taking shape!

Ichthyostega 3D print taking shape!

Then, on Saturday for our second free event we join forces with Ben Garrod (master of primate evolution, the secrets of bones, and “Attenborough and the Giant Dinosaur”) and RVC’s forensic pathologist Alexander Stoll as well as Sophie Regnault (“sesamoid street” PhD student w/me). As the “Large Animal Dissection” title hints, it’s not the right kind of gig to bring small kids to. There will be blood and stuff— we’ll be dissecting a cheetah together from 10am-4pm. This will involve walking through all the major organ systems, giving evolutionary anecdotes, and plenty more, with an aim to understand how the magnificent adaptations of cheetahs evolved—but also to investigate what problem(s) this animal faced that led to its sad demise. By the day’s end, there will just be a skeleton left. Get a front row seat early for this event, which serendipitously ties into “Team Cat”’s Leverhulme Trust-funded research project (we wanted a big animal and it just happened to be a cheetah; I had hoped for a giant croc or a shark or something but can’t complain!).

Ichthyostega 3D print is ready!

Ichthyostega 3D print is ready!

If you miss these events, please do cry bitter tears of regret. But don’t despair, there will be another “big cat dissection” in the London area in ~November (watch here for details), and plenty more fossil tetrapod stuff to come, plus a LOT more dinosaurs on the horizon!

Guess the bones! (photo by Zoe Self)

Guess the bones! (photo by Zoe Self)

And please come back to this blog post for more pics and stories as the week carries on… For hashtag afficionados, you can follow the fun on Twitter etc. at #firststepsCSF16. What a world we live in!

Update 1: While you’re here, check out our Youtube playlists of tetrapod-related videos:

Lobe-finned fishes

Ichthyostega‘s awesome anatomy

Tetrapod evolution: Tiktaalik to salamanders!

Update 2: Photos of our main stand (about tetrapod evolution)


Our poster/banner display looks nice.


Our tent brings in some punters.


Our bones excite people young and old, sighted and blind.


Fun with stickers and lab t-shirts.


Update 3: Cheetah meat & greet
Ben, Alex, Sophie and I tackled the cheetah dissection today and it went GREAT! Much better than I’d optimistically expected. Rain didn’t scare the crowds off and neither did the gore, which there was some of (gelatinous spinal cords, lumpy tumors and at least one flying tiny bit of cheetah flesh that landed on a good-natured audience member!). Photos will tell the tale:




Sophie and Alex help us get set up in our tent.


Our initial rough schedule- although we ended up improvising more after lunch.


Dissectors assemble!


The beast revealed. It was skinned by the museum that loaned it to us.


Alex showing his talent: removing the viscera in one piece from end to end, starting with the tongue.


Impressive finding of a surgical fixture (plate and wires) on the tibia, which had been used to hold the shattered bone back together long enough for it to heal. Added to the kidney disease and liver-spleen-lung cancer, this cheetah was in the sorriest shape of any cadaver I’ve seen yet.


Cheetah coming to pieces: (from bottom) lumbar/pelvic region, hindlimb, thorax, forelimb and other bits.


Dr Adam Rutherford, an eye expert, did a nice dissection of the cheetah’s eye, here showing the tapetum lucidum (reflective membrane), which shows up as light blue colour. Its small size befits the not-very-nocturnal habits of cheetahs.


The lens of the cheetah’s eye. Now cloudy because of dehydration and crystalization, but still fascinating to see.

Want to see more images and the enthusiastic responses from the audience (we got some great feedback)? Check out Twitter’s #cheltscifest feed, or more simply my Storify condensation of the cheetah-related tweets here.

I hadn’t been feeling very well for several weeks and then last night it happened. This post is a description of what it’s like to be an epileptic, written simply to document my experience. My goal here is to do that, almost in a dispassionate scientific way, and if it helps others going through similar experiences — feedback I often receive from such posts — that’s wonderful. My post is not a call for sympathy or help, although those are understandable and kind responses, and it’s not a complaint either. It just is, because what I am is what it is.

I’ve realized that my blog has become about not just documenting how amazing, freakish and immensely fallible that anatomy can be in other species, but also about my own experiences with my anatomy (and physiology) failing, as per these two prior posts about my shoulder and brain (more links therein). Sharing these experiences gives me strength and clarity, even if some of that emerges from partly detaching myself from the emotional nature of the experience and trying to look at it from outside of myself. I can be a private person, so feeling like I can discuss something uncomfortable and vulnerable makes me feel like I am growing, much as I innately resist that.

Stomach-Churning Rating: 1/10; no fun events described, but no images either.

I’ve had enough experience now as an epileptic that I look back on my seizures with disappointment (“Oh damn, not again.”) but also familiarity (“OK that happened; I know how things will go now.”). They are terrifying at the time, especially for my family, and my disorientation when emerging from unconsciousness with strangers around and with a gap in my memory is nightmarish.

I was watching a documentary about the Jutland battle in WWI while my daughter was put to bed. Then… I woke up, maybe 20 minutes later, unsure what was going on. There were two “first responders” (emergency non-paramedics) present, one of whom I eventually recognized from my prior emergency experiences in recent months, trying to talk to me with my wife. I was impressed to later hear that they’d come within 5 minutes of being called; not bad for life in a small English town. I came to realize that my right shoulder hurt again (from violent spasms), reducing it to almost a one-degree-of-freedom joint (mostly able to move fore and aft; almost zero pronation/supination without intense pain), reminscent of Ichthyostega‘s. I was surrounded by tissues wet with blood from my lip, where I’d again bitten myself during my fit. I could sense my racing heartbeat and fluctuating temperature, other hallmarks of my pre- and post-seizure symptoms. My vision was blurry, with my eyes usually becoming dilated during a seizure.

But the predominant feeling that takes an uncomfortably long time to pass is the “post-ictal state“, a mind clouded by confusion, slowly becoming aware that my neurons are misfiring but are beginning to sort themselves out. I sometimes irrationally want to just go back to sleep and not talk, and need some rational insistence from carers that we can’t do that right now. It is this vacillation between consciousness and unconsciousness, in a grey area in between, that I find most disturbing, as I cannot completely trust my own mind, disbelieving what is happening (“Is this real?”), and sometimes I lapse back into seizure(s) again. This is a powerful example of the frightfulness of uncertainty. As a scientist, so reliant on my mind, it is horrifying to feel like it is out of control. It also conjures up memories of observing my mother’s mind declining with Alzheimer’s syndrome, and those are vastly painful.

As I became able to put words together semi-coherently, and as the medics poked and prodded me to do tests on my symptoms (I had a cannula in my left arm’s blood vessel by now), discussion turned to whether to take me to the hospital once the paramedics arrived with the ambulance. In the past, there was no question of the need for a trip to Accident & Emergency (A&E in the UK; same as the ER in the USA).

Yet now, with almost 2 years of experiences behind me, I (and my carers) have come to know my better seizures from my worst ones. And given that A&E normally involves >4 hours of lying around in a noisy room, constantly disturbed by checkups or screaming patients, it is far from restful and rest is what I tend to need most. After an hour of vigilance, my symptoms faded and I became more able to answer queries, even to talk over options. We agreed that I could stay home, try to rest, and go to A&E if I had another seizure.

I am glad to say that I got a full night of rest and I feel a lot better today. That I am able to think clearly enough to write this post gives me reassurance. After past seizures, I’d often be unable to do much except take naps and gawk slack-jawed at the TV screen, with my vision still blurred (one eye even seemed to change shape post-seizure once, and I began seeing things in the corner of my field of view that are not truly there). So the bright side of this post is, maybe my medications are working better now, and maybe we will get this epilepsy under control, but I keep saying that every 2-3 months and then being proven wrong by another seizure, so a lot of uncertainty looms.

Nonetheless, seizures involve a “refractory period” that makes further seizures less likely for some time period, so odds are good that I can feel more secure while I recover from this event, which usually takes two weeks or so to get my brain feeling closer to “normal”. Even so, my mind remains clouded by these post-ictal feelings, weighting me down with fatigue that is the most chronic challenge I struggle against now as an epileptic. It leaves me unable to do as much as I once could, with a backlog of work growing behind me like never before. This is the “diminishing” that I lamented before; it is not just old age.

That’s what one experience was like for me, and I’m glad that it was far from the worst “neural storms” I’ve suffered. I hope that readers find it interesting. Now that my battle is over for now, I’ll take some time to find out how that Jutland battle turned out.

Last year we finally, after about 14 years of slow work, released our biomechanical model of an ostrich’s hindleg. We showed how it informed us about the potential leverages (moment arms; contributions to mechanical advantage of the joints) of all of the muscles. It was a satisfying moment, to understate it, to finally publish this work from my postdoc at Stanford. Today, we begin to deliver on that model’s promise. And it only took 4 years or so, roughly? The journal Royal Society Interface has published our study of how we used this musculoskeletal model to simulate walking and running dynamics. Those simulations join an intimidatingly broad and complex literature using similar models to study human (and some other primate) locomotion or other functions at the level of individual muscles (for whole limbs/bodies) in vast detail and growing rigor. I have Dr. Jeffery Rankin, a research fellow finishing up his post with me after ~6 years of hard work on many projects, to thank for driving this work forward, and Dr. Jonas Rubenson (now at PennState) for his patient collaboration that has continued since the early 2000’s.

Stomach-Churning Rating: 2/10; computer models of muscle actions. The underlying anatomical data are goopy, as prior ostrich-dissection-focused posts show!

Our model; in right side view (on the left) and frontal view (on the right), with muscles in red and the leg's force as the blue arrow; frozen at the middle of a step.

Our model; in right side view (on the left) and frontal view (on the right), with muscles in red and the leg’s force as the blue arrow; frozen at the middle of a running step.

Simulations like these predict things that we can’t easily measure in living animals, such as how much force muscles and tendons generate, how quickly those tissues change length, how much mechanical energy they thus contribute to the joints, limbs and whole body, how much metabolic energy their actions cost, and much more. There are more ways to use these tools than I have space or time to explain, but simply put we forced our ostrich simulation to match experimental measurements of the motions and forces of a representative walking and running ostrich stride, from contact of one foot until the next time that foot hit the ground. It then used optimization methods (minimizing target criteria like muscle stress) to estimate how the muscles and tendons were used to generate those motions and forces. This is a ways ahead of some prior ostrich simulations such as this one that I recall from classes during my PhD studies.

Any modeller worth their salt knows that their models and simulations are wrong at some level. This is much as most science is “wrong”; i.e. a simplification of reality with some errors/noise introduced by assumptions, variation, methods and such. But generally these kinds of musculoskeletal dynamic simulations hold up pretty well against experimental data. A standard “validation” is to test how closely the simulations’ predictions of muscle activity match the “real” (measured in life, also with some uncertain error) activity of muscles. Science still lacks those data for ostriches, but fortunately measurements from other birds (by Steve Gatesy and colleagues) indicate that muscles tend to follow fairly conservative patterns. Grossly speaking, avian leg muscles tend to either be active mainly when the foot is on the ground (stance phase) or off the ground (swing phase). Some studies acknowledge that this is an oversimplification and other muscles do act across those two phases of a stride, either in multiple pulses or as “transitional” (stance-to-swing or swing-to-stance) switch-hitters in their activations. Our ostrich predictions matched the qualitative patterns for avian muscle activations measured to date, so that’s good. The results also reinforced the notion of transitional or multi-phasic muscle activation as still having some importance, which bears more study.

Yet what did the simulations with our ostrich model tell us that other ostrich experiments or other bird species didn’t? Three main things. First, we could calculate what the primary functions of muscles were; they can act as motors (generating energy), brakes (absorbing energy), springs (bouncing energy back and forth) or struts (just transmitting force). We could then sum up what whole muscle groups were doing overall. The image below shows how these broad functions of groups vary across the stance phase (swing phase is harder to condense here so I’ve left it out).

Positive work can speed things up; negative work can slow things down.

Positive work can speed things up; negative work can slow things down. Solid bars are running; striped bars are walking. (from our Fig. 13) You may need to click to em-broaden this image for the gory biomechanical details.

There’s a lot going on there but a few highlights from that plot are that the hip extensor (antigravity) muscles (biarticular hip/knee “hamstrings”) are acting like motors, the knee extensors (like our quadriceps) are mainly braking as in other animals and the ankle is fairly springy as its tendons (e.g. digital flexors; gastrocnemius) suggest. We often characterize birds as “knee-driven” but it’s more accurate biomechanically to say that their hips drive (power; i.e. act as motors) their motion, whereas their knees still act as brakes — in both cases as in many other land vertebrates. Thus in some ways bird legs don’t work so unusually. Birds like ostriches are, though, a little odd in how much they rely on their hamstring muscles to power locomotion (at the hip) rather than their caudofemoral muscles, which are reduced. Zooming in on some particular muscles such as parts of the hip or knee extensors, the functions sometimes weren’t as predictable as their similar anatomy might suggest. Some muscles had parts that turned on during swing phase and other parts used during stance phase. Neural control and mechanics can produce some unexpected patterns.

Second, we looked at one important methodological issue. Simulations of musculoskeletal dynamics can vary from simple static (assuming each instant of a motion is independent from the others; e.g. ignoring acceleration, inertia, tendon effects, etc.) to more complex grades of fully dynamic flavours (e.g. assuming rigid or flexible tendons). We looked across this spectrum of assumptions, for both walking and running gaits, with the expectation that more static assumptions would work less well (vs. more dynamic ones; by various criteria) for running vs. walking. This also showed us how much tendons influence our simulations’ estimates of muscle mechanics—a fully rigid tendon will make the muscle do all of the work (force times length change) whereas a flexible tendon can literally take up some (or even all) of that slack, allowing muscles to remain closer to their isometric (high force-generating, negligible length change) quasi-optimum.

Nicely, our predicted muscle functions weren’t influenced much by these methodological variations. However, static assumptions  clearly were in some ways less appropriate for running than for walking, as were flexible tendons. Somewhat surprisingly, making the simulations more dynamic didn’t lower the levels of activation (and thus presumably the metabolic costs) of muscles, but actually raised those levels. There are good reasons why this might be realistic but it needs further study. It does muddy the waters for the issue of whether assuming that rapid locomotion can be modelled as static is a “bad” thing such as for estimating maximal speeds—yes, tendons can do more (elastic energy storage, etc.) if more dynamic models are used, but co-contraction of antagonistic muscles against each other also brings in some added costs and might lead to slower speed estimates. We’ll see in future work…

Finally, one often overlooked (sometimes even undiscussed!) aspect of these simulations is that they may silently add in extra forces to help muscles that are struggling to support and move their joints. The justification is typically that these extra “reserve actuators” are passive tissues, bony articular forces and other non-muscular interactions. We found that the hip joint muscles of ostriches were very weak at resisting abduction (drawing the thigh away from the body) and this needed resisting during the stance phase, so our simulations had very high reserve actuators switched on there. That fits the anatomy pretty well and needs more investigation.

Want to know more? Happily, not only is the paper free for anyone to view but so are all of the data including the models (modified slightly from our last paper’s). So, although the software (Opensim) isn’t ideal for 4-year-olds to play with (it is fancy engineering stuff), if you have the interest and dilligence it is there to play with and re-use and all that. But also watch this space for future developments, as there is more to happen with our steadily improving models of ostriches and other beasties. Anyway, while this paper is very technical and challenging to explain I am not too bashful to say it’s one of the finer papers from my career; a big stride forward from what we’ve done before. I have been looking forward for a long time to us getting this paper out.

P.S. Our peer reviewers were splendid- tough but constructive and fair. The paper got a lot better thanks to them.

I’m now asked all the time how I’m holding up in light of recent changes in my life, and I have a hard time answering that question, as I am still not sure – except that I am still here, more or less. Over the past year I’ve grown to embrace the notion, reinforced very strongly by my own frequent experiences, that I am disabled (in the medical/legal sense). I’ve had a harder time embracing the idea that I am now part of a minority group (I have strongly accepted that I am a senior, white, male scientist in the upper-middle class; which conflicts with “minority” in every way). Categorization aside, one conclusion I’ve been grappling with is the strong sense that I have been diminished. I can’t think as clearly, my memories are fading and my body is increasingly decrepit in physical and physiological ways that are becoming obvious to me. And yet I struggle to explain these feelings to people. So here I am, writing a blog post that is partly about what it’s like to feel that my personal “glory days” as a human scientist have passed. It’s not very uplifting stuff although there is a surprise of sorts at the end.

Stomach-Churning Rating: 1/10; a medical imaging scan of my disintegrating body, and a cartoon of surgery.

Before I go into my sob story, I should reinforce that there’s not just doom and gloom here. There is a heady, very complex mix of feelings. I’ve still got a great family (save the demise of all of my close relatives from my parental generation and before) and friends. Much of the time I’m still able to smile and have some fun. I’m not a Syrian refugee clinging to life in a tent while the world turns its back on me, or an Indonesian orangutan aloft in a tree watching the forest burn around it while the palm oil plantations spring up in the distance. All is relative. I know my life best and that’s what I blog about, so here is that infinitesimal perspective on life.

At work, I am buoyed by a fantastic team of scientists; some of the best I’ve ever worked with. They churn away at the science while I try to lead them. We’re doing some very hard science lately; some of the most challenging work I’ve been involved in. And my declined health hasn’t helped me to lead them, so sometimes they’ve had to rely on each other for spans of time. It’s not the best analogy but I often feel like we are in the open sea and I am swimming in front of our boat full of precious science, navigating while I barely keep my head above water and they struggle with the oars and their own exhaustion. In real life, it’s seeing their smiling faces and the wonderful science they show me on a regular basis that helps me keep afloat, personally.

Indeed, one take-home message of this post is that, while feel myself diminish, struggling with normal ageing and major new health problems, I see those I mentor grow and I get a vicarious thrill and pride from it. This is something that I know many research scientists experience to varying degrees, and often in conjunction with the too-often-metaphorized(?) experience of parenting and having the joy of seeing one’s offspring mature while one feels old age encroaching. As many research managers witness, I see my team’s collective research expand and build new levels of coolness in our little domain, I get wiser by reflecting on the successes and failures, and one could say I enjoy some credit from my team’s work by navigating our general course of research while they do the daily technical work and I help mentor them through their careers.

But first, a content cat.

But first, a content cat.

At the same time, as my collaborations and range of projects broaden, following my increasingly integrative interests, I see my relative expertise decreasing. It seems a long way now from my postdoc years, just over a decade ago, when I could run most or all of the software and hardware I needed to do the science. I am increasingly uncomfortable with that. However, I still learn new skills and knowledge so I am far from static as a scientist, and I am pushing myself more and more to learn more, wrestling with my age/health-imposed difficulties in learning. Furthermore, as a human being I feel far more aware of the world and the broader issues at stake than ever before (thanks in part to social media, I should add; but also thanks to my curiosity about life beyond my research). Improving my mind is still a goal of mine. The “diminishing” label I apply here is not that fair perhaps, but it’s how I feel about myself and I am sure there is some truth to it. It is the foe I grapple with.

So I get bemused reactions when I’m asked how it’s going and I respond, somewhat glumly, that I’m “hanging in there”. Many know I’m having health problems and tell me they are inspired by how I’ve held up and how well my (team’s) research and science communication and other work seems to be going, from the outside. Indeed, we’re cranking out more papers and surprising amounts of funding (see below) than ever before, so on paper it does look very good. It helps a little to hear those comments of how impressed some friends and colleagues are, but I don’t feel very impressed with myself. I feel lucky to have a great team of scientists and to have a great job in an insanely good laboratory environment, because otherwise things would be very different for me. I’m starkly aware of my privilege and feel vastly fortunate for having it.

My personal experience in work/life doesn’t reflect the joy of “success” that might seem to spring forth from my CV or the image that the outside world might get of me. I’m seesawing back and forth between those intermittent joys (and other happiness that comes from life away from work!) and a sense of hopelessness. I see the grim state of humanity and the broader world, and I look within and see my own decrepitude advancing, and I feel sad. It’s not a clinically manic-depressive seesaw but I can see some similarities when I apply my scientific detachment skills and look at myself from a quasi-objective perspective. I’m not the naïve, “everything is excellent” optimistic grad student I was—I just see flickers of that person these days. Sometimes I like to see him.

Much of the torques applied to the seesaw come from my oscillating health status- I alternate between good and bad days, with hints of a broader weekly rhythm that my physicians and I are still trying to grasp. Those oscillations determine everything for me: I can be bursting with energy and make strong inroads on my “to do” list, or I can be utterly drained and unable to do much more than stare vacantly or maybe fire off some emails to make incremental progress on work. I tend to be lingering somewhere in the middle, with far less vitality on average than I had two or so years ago. I look back on those past years and feel like I am looking up at the peak of my life and career. Time will tell if that’s “true” in some way or not.

Regardless, over the past 18 months there have been huge “valleys” from when I end up in hospital after major epileptic attacks, with a couple of weeks of recovery afterwards. Overall, my capacity to do what I used to be able to do has been halved. At best, I feel like I can operate at maybe 90% of my peak capacity and that never lasts long. Some of this is the inevitable decline that comes with entering one’s mid-forties, but some is a new step-change that has hit me over the past 18+ months since I became an epileptic suffering from tonic-clonic (“grand mal”) seizures every 2-3 months. Why is this suddenly happening and why haven’t the doctors resolved it yet? Well, the short answers are that my brain had damage (I told that story here) that can lead to epilepsy later in life, and that medicine still isn’t perfect. Epilepsy that cannot be entirely suppressed by existing medication is still common. We’re still experimenting with medications for me but it’s too soon to tell if we’ve found a solution, and we might never.

I heard some wise words a while ago that “we’re more content to blame ourselves than to accept that some things are beyond our control” and I’ve taken that to heart. Life is scary and short and it’s true that a lot is out of our control, especially the end of life. In reflection, I’ve been tempted to look back on choices I made in life and try to blame myself for what damage that has wrought on me (or others), but in terms of health I question that assumption. I may just be the victim of bad luck (genes, etc), but some people find bad luck too hard to accept, implying an indifferent universe rather than free will leading to misfortune/fortune. I’m not out of hope but I’ve accepted that the current state of my life might be just how it will be, and that’s been a hard lesson, but one I’ve learned again and again with my many chronic health problems over 20+ years. I don’t blame myself (much) for all that. More than ever, I appreciate the other wise words that “everyone is fighting a struggle you know nothing about”. I might look to an external observer like I’m kicking ass, but I feel anything but that kind of triumphant, fist-pumping jubilation.

I feel lucky to still be here, and eager to keep it that way, but I am so, so tired. Intellectually, physically, emotionally, it’s like a vampire has been paying me regular visits. And so I have to sigh, more than I used to, when confronted with bullshit like excessive paperwork or petty politics or something else I wish I didn’t have to endure, deeply feeling life slipping past as I do endure it, but that’s life for you. And at work, as a senior research manager, that’s often my job to endure it, in ways I’d never experienced as a junior researcher. I just have to cope with being pummelled by waves of difficulties and not grow weaker if I can avoid it. Coupled with life’s other burdens, the diminishing scientist faces a different beast of challenges and can often feel very alone. It’s a strange new place I’ve found myself in, far more complex than the worries I had as a postdoc, with harder choices to make and vast grey areas to traverse.

Nonetheless, as welfare science likes to term it, it’s entirely “a life worth living”. I have to pick my battles more than I used to, and I’ve had to learn to take more time to get exercise, rest, and avoid the stresses (or even unpleasant people) that can cause my health to take rapid downward spirals. I’m more fragile in many ways, such as having to stop doing karate because my shoulders have weakened. Here’s some interesting anatomy for you from a recent MRI scan of my right shoulder:

My left shoulder in top cross-sectional view, with the missing parts of my humeral head crudely outlined in red. There's more amiss here, too.

My left shoulder in top cross-sectional view, with the missing parts of my humeral head crudely outlined in red. There’s more amiss here, too.

My seizures cause my shoulder flexors to spasm, raising my arms up and crushing my humerus against my glenoid cavity of my scapula and causing occasional dislocations that abrade the humerus against the rim of the glenoid. The result, after numerous seizures, has been the wearing away of the articular cartilage of my shoulder and then the crumbling of the bony head of my humerus. Thus, once my NHS surgeon is ready to in coming months, I am due to have my coracoid process of my scapula cut off and moved, with its attached muscles and ligaments, to be screwed into the front of my glenoid cavity, bracing my humeral head more tightly against the glenoid and thereby resisting future dislocations. Luckily that operation can be done with several small incisions and endoscopy; invasive as the surgery is; thus recovery time won’t be so long.

Latarjet surgery (view of right shoulder joint [glenoid] from front): coracoid process moved posteroventrally. More details (w/videos) here.

Latarjet surgery (view of right shoulder joint [glenoid] from front): coracoid process moved posteroventrally. More details (w/videos) here.

It amuses me that all of this intense surgery looming on the horizon doesn’t worry me. I just want it done. I’ve been through a comparable surgery with my left shoulder, involving screwing my greater tuberosity back onto my humerus, so I know what recovery is like, and now that shoulder is doing fine. All that aside, my physical integrity has declined and I feel it every day. I may never return to my karate classes and earn that black belt I was seeking as a life-goal, but time will tell. I am trying to do what I can to remain as strong as I can for as long as I can.

A year from now all of my major funding and most or all of my research team were due to finish or be finishing. Over the past year, I was thinking forward to this eventuality and truly looking forward to having a smaller, quieter team, with less pressures on me. Many of those pressures are self-imposed because I am still ambitious and love doing science. I can still feel that youthful passion welling up inside me sometimes, so strong that I imagine it to be a tidal wave that could consume the world. It fuels my drive to try to do more, better science, but is dampened now by the problems I’ve lamented above, but it’s still there. So that passion and drive led me to, on a whim, resubmit an EU grant that was rejected a couple of years ago. I didn’t take it super seriously and so writing the grant didn’t stress me out. But a week after submitting it, I was back in hospital anyway, in bad shape. Over the following nine months, I grew to hope that the grant wasn’t awarded and expected that it wouldn’t (given <20% funding rate especially as a young Advanced Investigator in that ERC funding programme; https://goo.gl/Ps0Rhd if you want to know what that means). A big part of me still wanted to have that smaller team and less (or no) funding. I’d even contemplated leaving academia. I dream sometimes of retiring early to a quiet life with my family or wandering off into some jungle for a foolish adventure, but neither is realistic.

Yet a few weeks ago, the email from the EU came with an answer. I got the grant: 2.5 million Euros for 5 years of research on dinosaur evolution and biomechanics. More about that later. The funding details are still in negotiation but I now am on course to be advertising (in ~August) 4 new jobs to work with me for up to 5 years on this project, beginning in October. My reaction has puzzled those colleagues I’ve told about the grant, although I have kept that news quiet (until now) while I finish the paperwork for the grant award. I feel mixed about getting a large grant at this time in my life. It’s a helluva lot of work and five years seems a very, very long time to me, and to focus on one major theme—and to study dinosaurs.

I had also looked forward to moving away from dinosaur research—but, like Al says in the video above, their siren call can drag us back to the Mesozoic era with questions that entice us and with spectacular fossils that are a riot of fun to study. In this case, we’re going to be looking back on the “locomotor superiority” hypothesis that has been bounced around for >40 years as a possible explanation for why dinosaurs flourished whereas other archosaurs (except crocodylomorphs) didn’t, and how much bipedality relates to that, in terms of various behaviours and motions. Can these questions even be answered? We shall see.

Yes, boo hoo! Poor me, getting a coveted grant and all that! I am not surprised if that is hard for others to understand, and I still am figuring out how I feel about it all. Professionally, this is a wonderful thing; no question. Personally, it’s pressure I didn’t need to put my disabled, diminished self through. Irony and conundrum aside, I want to do it and I should try. Regardless, off I go, with a new-team-to-come and my research focus dominated by one main project, the largest grant I’ve ever managed (by a long shot!). It’s interesting times for me ahead. Life has come full circle, returning me back to science-ing the dinosaurs/archosaurs I’d focused on in my PhD work. But I am not the same person, and so it will be a very different experience. Somehow I have to balance this challenging project with the struggles in my life in general, and that will test me in diverse ways. I’m sure there will be many surprises in my work and personal life during the next five years, and I’ll be sharing them here on this blog when I can.

I’ve tried to express my own journey through the big ups and downs I’ve seen over two years. Maybe it will help others who are quietly, or noisily, struggling. I’m curious to hear from others that have experienced feelings of themselves declining as their careers/lives (in science/elsewhere) move along in some direction.


Goodbye Pedro (?/?/2014-23/4/2016). We had too little time together. What we shared was so lovely. Parting has been terrible sorrow.


I had a nice chat today (OK, a while ago, when I first started writing this post) with a researcher who wanted to know why I blogged and how I balanced my science communication work with my research activities. In talking with her, I realized that there was a way to explain my views that I hadn’t thought of before and then I realized that this might make a blog post that other people would care to read. So here it is.

Stomach-Churning Rating: 0/10. Just vomiting out words.

I’ll take a historical perspective to answering why I bother doing what I do, and how I came to do it.  As an evolutionary biologist, and as a teacher and communicator, I find that this kind of narrative works for me and for others, in deepening comprehension of why things are the way they are. In my pre-college and college years, I loved to read and write, and I liked science. A professor as a father, a mother who encouraged scholarship and a family that liked to go experience nature together helped. I eventually got competent at writing and maintained that ability (even musing about whether I might become a Hollywood screenwriter– bullet dodged?), whilst figuring out that some sort of biology might be right for my career. Off to college I went.

Amidst the beer-fuelled haze of undergrad life at the U Wisconsin, I fell in love with the science writings of Stephen Jay Gould and some others, which today would be blog posts. A course in the history of science and a lot of courses in biology shaped my interests, and to cut that story short, off to grad school I went in 1995. I’d discovered the internet (on my dad’s iMac) while recovering from some major health problems, and liked it. I learned to love it during my PhD work, participating in a lot of arguing on the Dinosaur email listserver and other kinds of online writing.

In particular, I fell in with the crew of grad students writing pages for the UCMP website— the original online virtual museum. These pages, intended as “virtual exhibits”, were more or less blog posts, or wikipedia-esque entries, or whatever you want to call them in modern parlance. They were reviews of and commentaries on current knowledge on various topics in biology/palaeontology. I now realize that even back then I was an avid science communicator, especially online. I now realize that this hasn’t truly changed in 20 years, except how I do that online communication.

I lacked the self-confidence (or experience) to do much science communication in person, preferring to take the time to slowly write posts/pages online while in hermitage in the museum basement’s computer lab or at our lab’s office computer. Oral presentations utterly terrified me (sometimes to the point of paralysis) until late in grad school once I’d had some practice doing them at seminars and conferences, then I started to love talking about science in person to larger groups (teaching undergrads helped build this love, too). My work began to attract the attention of the media, especially in 2001-2003 during my postdoc at Stanford once my T. rex and elephant biomechanics research got published, and that attention helped train me (through some great mentors’ help) to be better at reaching and engaging with the public via the media.

By 2003 I was a faculty member and my science communication activity continued, even expanding as I became more well-known. Yet my online “scicomm” presence was by this point reduced essentially to only my personal RVC webpages (with several stories about my research that were akin to blog posts) and occasional journalists’ stories about my research. I focused on being a “typical” scientist. That worked for me; I was happy with it.

Finally, in ~2010 I started to catch the online science communication bug again, inspired by bloggers and science writers (and scientists) like Darren Naish (Tetrapod Zoology) and informal chats with colleagues who were experimenting with the newer forms of communication, which were far more engaging than prior forms like the static, non-interactive webpages I’d worked on at the UCMP. And so here I am; see the “Welcome” tab for how this blog originated.

Why then do I continue with this and how to I find whatever balance exists between scicomm and research? First off, I don’t like to think too much about categorizing what I’m doing. It’s all science (writ large) to me, whether I am doing computer analysis or dissections or writing papers or training people or writing blog posts or tweeting. There is a “zero-sum game” that people obsess about in balancing scicomm “versus” research. Yes, it’s true: there are just 24 hours in a day, and to be a successful scientist one must spend some of those hours, on average, on research. Wow. But as long as the research is moving forward at a pace I’m happy with, I don’t care if it took me an average of 6.5 or 8 hours/day to get my part done, or how that time precisely was spent. I don’t think I ever have cared. I don’t do hours-accounting to ensure that I am clocking in and out the “correct” number of hours. I hate that shit and it’s why I didn’t go into a job where I must clock in and out. Lack of freedom suffocates me, and I think that the zero-sum mentality suffocates science.

Similarly, I don’t feel that some vital opportunity is lost by spending time on scicomm rather than research data. I’m sure that, over the years, I’ve “lost” a paper or two that I could have written instead of doing some blogging or other scicomm, but I have enjoyed the latter and others have too, so I feel that it has been more than worth whatever “cost” there has been. If some scientists feel that essentially all they should be doing in science/life is publishing peer-reviewed papers, that’s their opinion but it is not mine. But — I don’t consider that my blog posts are equivalent to peer-reviewed papers by any means, either. I list my blog on my CV as one line, not all of my blog posts like I do for my papers. It is far harder to publish a scientific paper than to write a blog post; there is no equivalency, at present.

Yet on the flip side, blogging is valuable and fills a gap that scientific papers cannot. Once I became a Professor in 2011, I felt even more liberated than when I got tenure. I felt that it was time for me to try new things and take risks, that this was what being a professor was partly about. So I looked around at opportunities and, reflecting on my experiences with documentaries like “Inside Nature’s Giants” and various conversations I’d had in person and on social media (mainly Twitter), I tried blogging. I saw opportunities to engage a broader audience in discussions about anatomy, and I wanted an outlet in which to be creative as a writer and scientist, so blogging fit me. In a blog like this, I can be human, I can talk about myself or others in a personal, less detached and dry way, I can make very speculative statements, I don’t have to reference everything or obsessively avoid any sloppiness, I can be casual, I can be longwinded (like this post), whatever — I can do what the fuck I please, including say “fuck”. I can write posts on anything I want and it doesn’t have to be a novel contribution nor do I have to “sell” it in ways that I might with papers. Try those things in a peer-reviewed publication. Research papers are straight-jacketed by rules because that constrained format has worked for them over the past two centuries. It works for transmitting carefully-checked results that assemble a body of increasingly-trusted knowledge.

I like the release that blogging gives me, to be intellectual in a more creative, explorative, personal, conversational way. This freedom is healthy and rewarding for me, I’ve found, giving me a satisfaction that I can’t get from just grinding away at papers, by engaging with the broader, (potentially) global public rather than just a few specialists — or just a few local people at public science events. Let’s face it, a lot of our peer-reviewed publications don’t matter much; they won’t get cited much and might eventually get forgotten, and even those few specialists that do care about them might not care that much. What matters to me is how I feel about how I am spending my life. I do passionately love discovering new things in my research but science doesn’t start and end there for me. Much like with social media, I’ve benefited from things (some unexpected) like new collaborations, meeting scientists/journalists/science-enthusiasts who become new friends, invitations to do new things like give seminars or take part in bigger media (documentaries, major websites, etc.), being seen as an expert by a broader audience, a stronger and more diverse CV (including positive comments on reviews for funding/other applications), improved writing/communication skills and more. Hence it would be false to say that I lose something by not spending all my time on conventional science. I gain other things from blogging and other scicomm that have their merits, and in the end only one thing matters: does it feel worth it to me? Yes. Do I think that everyone must/should do the same? No. To each their own, but more scientists should consider broadening their scicomm horizons, giving the potential benefits more thought in the weighing of all the priorities that they must juggle. Some people aren’t ready to add scicomm to their repertoire. Some just aren’t suited to it at all. That’s fine. It’s part of the diversity of science. Not everyone can be part of the same conversations and experiences.

To me (like so many in scicomm), whether it is papers or blog posts, it’s all about conversations. These formats of conversations have the same intent: to share and discuss. Indeed, they have much in common, formatting and rules aside. That’s why I like in-person interactions, or seminars and conferences, or social media, or other forms of conversation, too. Each has its pros and cons, and except for social media scientists have been doing them for centuries. One could consider social media/blogging to have been around for 20+ years depending on how one defines it. I’ve realized that I’ve been engaging in conversations over the internet in analogous ways for those decades, and that blogging is not that new for me after all. I see myself as taking a break from the UCMP webpage-writing in by 2001 to returning to public online scicomm in 2010-11, with Facebook acting as a midwife for that transition. Posting science stuff on Facebook to a few hundred friends (ten of whom might show signs of caring) just wasn’t enough for me by ~2011. Since then, I’ve found that the diversity of the world that I can engage with in all these conversations expands my horizons and teaches me new things about science (e.g. other fields I don’t know much about) and about people (e.g. other scientists/science-enthusiasts/journalists I hadn’t met), or even changes the way I look at my own research (e.g. major trends in open access, open data, etc.). This is another major benefit that I wouldn’t get from isolation in an ivory tower. All of these things are echo chambers, whether one is in isolation in the research field of evolutionary biomechanics or the larger area of scicomm that this blog explores, or scicomm on Twitter, or whatever. Mindfulness of that reality can help prevent one’s mind from becoming too full of those echoes and losing sight of the broader world, and I try to be mindful.

I’ve discovered that I like to share, that this is a big part of my personality. I had suppressed some of that side of myself in the early 2000s, developing trust issues that I see in many other scientists today. I imagined things like: if I share, what if someone criticizes me, or I say something wrong and get embarrassed, or someone steals my data/ideas, or someone doesn’t like how I seem to be spending my time, or something else goes wrong. These imagined risks came from stories I’d heard about naughty colleagues and other people, and things I’d experienced myself, some of which were very unpleasant (e.g. colleagues trying to use me, or taking my data or ideas; or people just being jerks). I focused on the negative aspects of sharing but didn’t thoughtfully weigh them against the positive. I now see what I had missed, although things worked out well for me anyway and I don’t deeply have regrets about what I didn’t do prior to 2010-11. I was more selfish, true, and that was strategic selfishness (focus on my research career) as well as irrational selfishness (avoid things that scared me as risky), and there are other things about me back then that I don’t like (e.g. more competitive, less helpful/collegial; which tends to come with secretiveness/lack of sharing) but I don’t beat myself up about them. I could have been much worse, and what matters now is the choices I make now, which I think about much more carefully than I did back then, with the benefit of wisdom gained through successes, failures and mistakes.

Similarly, I don’t like to harangue myself about whether I am doing “work” or “life” stuff. I’m not a fan of this dichotomy. It is an unfair attitude to force upon oneself, in my opinion. “Work vs. life” is set up to impose self-flagellation for doing the wrong thing at the wrong time. I believe in work-life integration, just as I believe in research-scicomm integration in science. I do what I need and want to do when it feels right to do those things. The ultimate goals are what matter most; I want a good life in all facets, including making the lives of those around me good, too. I seek happiness and enjoyment and satisfaction, and do what feels like it will best bring me those things at any moment, depending on how I feel. Sometimes I have energy inside that feels best directed toward writing a blog post. Sometimes I’m more excited about a paper. Or a movie, or a cuddle with cats, or whatever. Generally my #1 job priority (as a senior mentor to numerous people) is to help my team get its work done and published (see my prior posts on managing teams), and whatever other time remains when I feel like working is spent on other things, like my own research or scicomm or whatever. I try to avoid guilt-bludgeoning myself about whether I am working or “living” at 06:00 or 10:45 or on Sunday or Wednesday. But I do bother myself about ensuring I have time with my family and friends, which brings many flavours of happiness and enjoyment and satisfaction that science cannot. I don’t believe in working that much on weekends (but I do dabble sometimes, and I don’t overly guilt-trip myself if I do) and I do want to take time off after 5pm most nights with family, and I do need 7-8 hours of sleep or I’m extra grumpy and low on my science-fu levels. I’m happy with my work-life integration, and it seems to be playing out OK as long as I manage my stress levels and physical fitness.

In summary, to me blogging is part of the balance that I try to seek in life, and that balance is deeply personal, ever-changing, and enjoyable to me. There will never be a right answer to the question of what balance is “right” for any one person, but there is a threshold of contentment that can be reached in seeking that balance, and if one reaches that, then that’s good. There will be people that try to tell you that the balance you seek is wrong, and some of them will be worth listening to but others are best ignored or told to piss off. That doesn’t mean that you won’t have to convince people that your balance is “right” for you, whether it is your family or your peers or grant funding committees. I’m fortunate that experiences in terms of people criticizing my time spent blogging have been minimal for me (I know this is not true for plenty of others!), yet I feel well-prepared to defend my blogging interests if I’d have to.

But concurrently there is a question of trust: shouldn’t those parties evaluating your life-balance be accountable for trusting you to make your own decisions, and questioning whether the standards-of-balance they hold as ideal should apply to you or not? This will always be a source of some tension with some people—and maybe it should be, but it has to be, because we are not all the same people living the same life. Maybe it’s easier “just” to focus on cranking out data and papers and not doing much else in science, much like it might be (or seem) easier “just” to blog 100% of one’s time. Maybe there’s jealousy or insecurity or fear of change that wells up in those that do find a certain, seemingly simple, set of priorities to be the right balance for them in their life, but don’t like to see others seeking a different kind of balance. Keeping more balls in the air in juggling life’s balance ramps up the complexity and that can be more difficult to control; more unstable, even. All of these questions of priorities, trust and balance arise from our humanity and our diversity and present us with frequent choices about how to handle them. I’d like to end this post on a more positive note by suggesting that we celebrate these questions, and by doing so that we celebrate our diversity and humanity and become part of the dynamic kaleidoscope of science and the real world. Maybe we should worry less about judging how others balance their lives; lives whose details we probably know little about; and push ourselves to learn something from our fellow scientists and science enthusiasts by seeking to understand how they’ve arrived at the point in life that they’re at, and seeking to build edifices of trust that there are many ways to contribute to science. We might learn new things that could inspire us to change our own priorities, much as I did in my life journey that led to this blog’s inception. There is a lot of common ground to roam in discussions about balance in the lives of those passionate about science. Curiosity and trust are key components of that ground’s bedrock, like they are of science’s. Discovery and sharing are other parts, in which research and scicomm play important, interactive roles.

I’m curious to hear what others think of things I’ve raised in this post about why I’ve struck the current balance I’ve made in my life (particularly regarding blogging but similarly extensible to social media), and how that relates to the many ways that others can find balance, which might lead to more harmony between research and scicomm in science. Kumbaya?

Anatomy of an amoeba: why not? I like it.

Anatomy of an amoeba: why not? I like it.

Well I made it this far, and I might not have expected to last four years of blogging but I did last, and I’m very glad. Here is my customary retrospective with some “new” images. I’m glad to see that you’re still here, too: over 3200 blog subscribers (roughly doubled since 2014?) and some regular commenters remain– thank you! Like an undead horror, this blog keeps lurching forward, and it appreciates loyalty.


 Stomach-Churning Rating: 5/10- there’s a bit of blood and stuff. Nothing that unusual. If you’re like Rick (above; by /u/epicolllie), that won’t phase you by now.

The past year on the #JohnsFreezer blog felt quiet to me, and that’s largely because I was distracted by numerous things; if you know me (or follow my whining on Twitter), you probably can guess. But there were enough highlights to keep me feeling satisfied. Judging from your reactions, you liked my inside scoop on the T. rex Autopsy documentary that I consulted on, and a distant second place post was, hmm, that’s kinda surprising: my elephant foot dissection post. I’m rather pleased so many of you clicked on that one, actually. That’s a meat-and-potatoes post for this blog, much like my latest one on getting to know M. caudofemoralis (longus). But hey, check out the “Goat to be Seen” post if those are your kind of entries– it seems a lot of people missed that one, and it had a mix of quirkiness, unflinching raw anatomy, and art that still makes me smile.

I was browsing my photos and ran across this NHMUK exhibit of a small ungulate hooved limb vs. a nice honking big padded elephant foot. It elegantly gets across the biomechanical differences between these limb structures.

I was browsing my photos and ran across this NHMUK exhibit of a small ungulate hooved limb vs. a nice honking big padded elephant foot (both abstracted down to their fundamentals). It elegantly gets across the biomechanical differences between these limb structures. Bravo!

My rants about how sometimes it’s helpful for scientists to put the brakes on media coverage of their own research, and on “HONCOs” (honorary co-authors), also brought in the punters, as did the re-post about the not-so-bad aspects of self-promotion in science. Not so many people appeared to read the post about where ideas come from in science but it got a lot of tweets, which is a strange incongruity, yet my thoughts on how to manage a research team didn’t do any better (but if you read in between the lines, there’s a poignancy to that post). Anyway: good ole rants; ahh, it still feels good to have those off my chest, even after all these months. And writing them helps me sort out my own thoughts, if nothing else.

Hindlimb of a sea turtle that we dissected in 2015 after it came in for a clinical postmortem.

Hindlimb of a sea turtle that we dissected in 2015 after it came in for a clinical postmortem.

One of my greatest science heroes, “Neill” Alexander, got his due here, and there continue to be comments trickling in on that post from people who are reminiscing about his influence on their careers. That is definitely one of the posts on this blog that I feel best about, even after four years. It meant a lot to me, much as Neill has meant a lot to others. I also did an homage to museums, which in parallel (oddly, but enjoyably for me) became an(other) homage to avian kneecaps. I like them too, and museums of course, but they’re awesome in a very different way from Neill.

Team Cat is still cranking on our biomechanical and anatomical studies of felids- expect a lot of new stuff from us in 2016! Meanwhile, enjoy this spectacular taxidermy.

Team Cat is still cranking on our biomechanical and anatomical studies of felids- expect a lot of new stuff from us in 2016! Meanwhile, enjoy this spectacular taxidermy– and check out Dr. Andrew Cuff’s blog with the latest science and stories.

I learned a lot about my genome in this fourth year of blogging, and I delved into that with you, as part of my commitment to share what I learn about myself by poking around in my biology. Oh, and I just learned that the image depicting my genomic ancestry was this blog’s most-clicked image this year– that’s cool, and unexpected!

The past year was a big year for dinosaurs on this blog, with a post on the “Giant Dinosaurs of London” and another related to my cameo in the giant titanosaur documentary with Attenborough, but with a focus on dissecting dinosaurs, and a blatant bandwagon tribute to/musing on Jurassic World.

The very, very strange iguanodontian dinosaur Lurdusaurus (forelimb; note the big spiky thumb claw), which I was pleased to see at the natural history museum in Brussels, Belgium in 2015.

The very, very strange iguanodontian dinosaur Lurdusaurus (forelimb; note the big spiky thumb claw), which I was pleased to see at the natural history museum in Brussels, Belgium in 2015.

And finally, this blog had a baby, or a sister, or whatever, this year, and that has been a blast: Anatomy to You was born, thanks to Dr. Lauren Sumner-Rooney’s expert care and dedication to science communication. If you haven’t checked it out, now’s a good time, or offer to do a guest post for our “In Focus” section if you’ve got some anatomical science to share! Speaking of guest posts, Julia Molnar did a fabulous one about our paper on crocodile backbones this year, here on this blog.

I'm still cleaning up specimens from the freezers: here, some "emu butts" (tails) from a collaboration with Michael Pittman and Heinrich Mallison, and PhD student Luis Lama's past thesis work.

I’m still cleaning up specimens from the freezers: here, some “emu butts” (tails) from a collaboration with Michael Pittman and Heinrich Mallison, and PhD student Luis Lama’s past work. Something about these vertebrae fascinates me.

I didn’t deliver on some plans for this year, such as a komodo dragon anatomy post, but I did finally do the “better know a muscle” and “dissecting dinosaurs” posts I planned, and a few other things, so the year worked out well enough.

What’s coming in year 5 of this blog? I have no bloody idea; I have not gotten that far. I think we’ll all be surprised. Let’s make the most of it! (I will consider requests)

"We'll always have elaborate models of gorilla muscular anatomy in Paris."

“We’ll always have elaborate models of gorilla muscular anatomy in Paris.”



Happy Darwin Day from the frozen tundra sunny but muddy, frosty lands of England! I bring you limb muscles as peace offerings on this auspicious day. Lots of limb muscles. And a new theme for future blog posts to follow up on: starting off my “Better Know A Muscle” (nod to Stephen Colbert; alternative link) series. My BKAM series intends to walk through the evolutionary history of the coolest (skeletal/striated) muscles. Chuck Darwin would not enjoy the inevitable blood in this photo-tour, but hopefully he’d like the evolution. Off we go, in search of better knowledge via an evolutionary perspective!

There is, inarguably, no cooler muscle than M. caudofemoralis longus, or CFL for short. It includes the largest limb muscles of any land animal, and it’s a strange muscle that confused anatomists for many years– was it a muscle of the body (an axial or “extrinsic” limb muscle, directly related to the segmented vertebral column) or of the limbs (an “abaxial” muscle, developing with the other limb muscles from specific regions of the paraxial mesoderm/myotome, not branching off from the axial muscles)? Developmental biologists and anatomists answered that conclusively over the past century: the CFL is a limb muscle, not some muscle that lost its way from the vertebral column and ended up stranded on the hindlimb.

The CFL is also a muscle that we know a fair amount about in terms of its fossil record and function, as you may know if you’re a dinosaur fan, and as I will quickly review later. We know enough about it that we can even dare to speculate if organisms on other planets would have it. Well, sort of…

Stomach-Churning Rating: 8/10. Lots of meaty, bloody, gooey goodness, on and on, for numerous species. This is an anatomy post for those with an appetite for raw morphology.

Let’s start from a strong (and non-gooey) vantage point, to which we shall return. The CFL in crocodiles and most other groups is (and long was) a large muscle extending from much of the front half or so of the tail to the back of the femur (thigh bone), as shown here:

Julia Molnar's fabulous illustration of Alligator's limb muscles, from our 2014 paper in Journal of Anatomy.

Julia Molnar’s fabulous illustration of Alligator‘s limb muscles, from our 2014 paper in Journal of Anatomy. Note the CFL in blue at the bottom right.

As the drawing shows, the CFL has a friend: the CFB. The CFB is a shorter, stumpier version of the CFL restricted to the tail’s base, near the hip. The “B” in its name means “brevis”, or runty. It gets much less respect than its friend the CFL. Pity the poor CFB.

But look closer at the CFL in the drawing above and you’ll see a thin blue tendon extending past the knee to the outer side of the lower leg. This is the famed(?) “tendon of Sutton“, or secondary tendon of the CFL. So the CFL has two insertions, one on the femur and one (indirectly) onto the shank. More about that later.

Together, we can talk about these two muscles (CFL and CFB) as the caudofemoralis (CF) group, and the name is nice because it describes how they run from the tail (“caudo”) to the femur (“femoralis”). Mammal anatomists were late to this party and gave mammal muscles stupidly unhelpful names like “gluteus” or “vastus” or “babalooey”. Thanks.

But enough abstract drawings, even if they rock, and enough nomenclature. Here is the whopping big CFL muscle of a real crocodile:

Huge Nile crocodile, but a relatively small CFL.

Huge Nile crocodile, but a relatively small CFL.

Bigger crocs have smaller legs and muscles.

Bigger crocs have smaller legs and thus smaller leg muscles, relatively speaking. CFL at the top, curving to the left.

The giant Nile croc's CFL muscle removed for measurements.

The giant Nile croc’s CFL muscle removed for measurements. 2.35 kg of muscle! Not shabby for a 278 kg animal.

However, maybe crocodile and other archosaur CFL muscles are not “average” for leggy vertebrates? We can’t tell unless we take an evolutionary tack to the question.

Where did the CFL come from, you may ask? Ahh, that is shrouded in the fin-limb transition‘s mysteries. Living amphibians such as salamanders have at least one CF muscle, so a clear predecessor to the CFL (and maybe CFB) was present before reptiles scampered onto the scene.

But going further back through the CF muscles’ history, into lobe-finned fish, becomes very hard because those fish (today) have so few fin muscles that, in our distant fishy ancestors, would have given rise eventually to the CF and other muscle groups. With many land animals having 30+ hindlimb muscles, and fish having 2-8 or so, there obviously was an increase in the number of muscles as limbs evolved from fins. And because a limb has to do lots of difficult three-dimensional things on land while coping with gravity, more muscles to enable that complex control surely were needed.

OK, so there were CF muscles early in tetrapod history, presumably, anchored on that big, round fleshy tail that they evolved from their thin, finned fishy one — but what happened next? Lizards give us some clues, and their CFL muscles aren’t all that different from crocodiles, so the CFL’s massive size and secondary “tendon of Sutton” seems to be a reptile thing, at least.

Courtesy of Emma Schachner, a large varanid lizard's very freshly preserved CFL and other hindlimb muscles.

Courtesy of Emma Schachner, a large varanid lizard’s very freshly preserved CFL and other hindlimb muscles.

Courtesy of Emma Schachner, zoomed in on the tendons and insertions of the CFL muscle and others.

Courtesy of Emma Schachner, zoomed in on the tendons and insertions of the CFL muscle and others. Beautiful anatomy there!

Looking up at the belly of a basilisk lizard and its dissected right leg, with the end of the CFL labelled.

Looking up at the belly of a basilisk lizard and its dissected right leg, with the end of the CFL labelled. It’s not ideally dissected here, but it is present.

An unspecified iguanid(?) lizard, probably a juvenile Iguana iguana, dissected and showing its CFL muscle at its end. The muscle would extemd about halfway down the tail, though.

An unspecified iguanid(?) lizard, probably a juvenile Iguana iguana, dissected to reveal its CFL muscle near its attachment to the femur. The muscle would extend further, about halfway down the tail, though.

Let’s return to crocodiles, for one because they are so flippin’ cool, and for another because they give a segue into archosaurs, especially dinosaurs, and thence birds:

A moderate-sized (45kg) Nile crocodile with its CFL muscle proudly displayed.

A moderate-sized (45kg) Nile crocodile with its CFL muscle proudly displayed. Note the healthy sheath of fat (cut here) around the CFL.

American alligator's CFL dominates the photo. Photo by Vivian Allen.

American alligator’s CFL dominates the photo [by Vivian Allen].

Black caiman, Melanosuchus, showing off its CFL muscle (pink "steak" in the middle of the tail near the leg).

Black caiman, Melanosuchus, showing off its CFL muscle (pink “steak” in the middle of the tail near the leg), underneath all that dark armour and fatty superficial musculature.

A closer look at the black caiman's thigh and CFL muscle.

A closer look at the black caiman’s thigh and CFL muscle.

Like I hinted above, crocodiles (and the anatomy of the CFL they share with lizards and some other tetrapods) open a window into the evolution of unusual tail-to-thigh muscles and locomotor behaviours in tetrapod vertebrates.

Thanks in large part to Steve Gatesy’s groundbreaking work in the 1990s on the CFL muscle, we understand now how it works in living reptiles like crocodiles. It mainly serves to retract the femur (extend the hip joint), drawing the leg backwards. This also helps support the weight of the animal while the foot is on the ground, and power the animal forwards. So we call the CFL a “stance phase muscle”, referring to how it mainly plays a role during ground contact and resisting gravity, rather than swinging the leg forwards (protracting the limb; i.e. as a “swing phase muscle”).

The “tendon of Sutton” probably helps to begin retracting the shank once the thigh has moved forward enough, facilitating the switch from stance to swing phase, but someone really needs to study that question more someday.

And thanks again to that same body of work by Gatesy (and some others too), we also understand how the CFL’s anatomy relates to the underlying anatomy of the skeleton. There is a large space for the CFL to originate from on the bottom of the tail vertebrae, and a honking big crest (the fourth trochanter) on the femur in most reptiles that serves as the major attachment point, from which the thin “tendon of Sutton” extends down past the knee.

Femur bones (left side) from an adult ostrich (Left) and Nile crocodile (Right).

Femur bones (left side; rear view) from an adult ostrich (left) and Nile crocodile (right). Appropriate scale bar is appropriate. The fourth trochanter for the CFL is visible in the crocodile almost midway down the femur. Little is left of it in the ostrich but there is a bumpy little muscle scar in almost the same region as the fourth trochanter, and this is where the same muscle (often called the CFC; but it is basically just a small CFL) attaches.

That relationship of the CFL’s muscular anatomy and the underlying skeleton’s anatomy helps us a lot! Now we can begin to look at extinct relatives of crocodiles; members of the archosaur group that includes dinosaurs (which today we consider to include birds, too), and things get even more interesting! The “tendon of Sutton”, hinted at by a “pendant” part of the fourth trochanter that points down toward the knee, seems to go away multiple times within dinosaurs. Bye bye! Then plenty more happens:

A large duckbill dinosaur's left leg, with a red line drawn in showing roughly where the CFL would be running, to end up at the fourth trochanter. Many Mesozoic dinosaurs have skeletal anatomy that indicates a similar CFL muscle.

A large duckbill dinosaur’s left leg, with a red line drawn in showing roughly where the CFL would be running, to end up at the fourth trochanter. Many Mesozoic dinosaurs have skeletal anatomy that indicates a similar CFL muscle.

We can even go so far as to reconstruct the 3D anatomy of the CFL in a dinosaur such as T. rex ("Sue" specimen here; from Julia Molnar's awesome illustration in our 2011 paper), with a fair degree of confidence.

We can even go so far as to reconstruct the 3D anatomy of the CFL in a dinosaur such as T. rex (“Sue” specimen here; from Julia Molnar’s awesome illustration as part of our 2011 paper), with a fair degree of confidence. >180kg steak, anyone?

As we approach birds along the dinosaur lineage, the tail gets smaller and so does the fourth trochanter and thus so must the CFL muscle, until we’re left with just a little flap of muscle, at best. In concert, the hindlimbs get more crouched, the forelimbs get larger, flight evolves and voila! An explosion of modern bird species!

Ozburt (72)

Left femur of an ostrich in side view (hip is toward the right side) showing many muscles that attach around the knee (on the left), then the thin strap of CF muscle (barely visible; 2nd from the right) clinging near the midshaft of the femur.

Another adult ostrich's CF muscle complex, removed for study.

Another adult ostrich’s CF muscle complex, removed for study. Not enough ostrich myology for you yet? Plenty more in this old post! Or this one! Or this one… hey maybe I need to write less about ostriches? The CF muscle complex looks beefy but it’s no bigger than any other of the main hindlimb muscles, unlike the CFL in a crocodile or lizard, which puts everything else to shame!

STILL not enough ostrich for you yet? Take a tour of the major hindlimb muscles in this video:

And check out the limited mobility of the hip joint/femur here. No need for much femur motion when you’re not using your hip muscles as much to drive you forwards:

But I must move on… to the remainder of avian diversity! In just a few photos… Although the CF muscles are lost in numerous bird species, they tend to hang around and just remain a long, thin, unprepossessing muscle:

Chicken's right leg in side view. CFC (equivalent of CFL) muscle outlined and labelled.

Chicken’s right leg in side view. CFC muscle (equivalent of CFL; the ancestral CFB is confusingly called the CFP in birds, as it entirely resides on the pelvis) outlined and labelled.

A jay (species?) dissected to show some of the major leg muscles, including the CF. Photo by Vivian Allen.

A jay (species? I forget) dissected to show some of the major leg muscles, including the CFL-equivalent muscle; again, smallish. [Photo by Vivian Allen]

Finally, what’s up with mammals‘ tail-to-thigh CF-y muscles? Not much. Again, as in birds: smaller tail and/or femur, smaller CF muscles. Mammals instead depend more on their hamstring and gluteal muscles to support and propel themselves forward.

But many mammals do still have something that is either called the M. caudofemoralis or is likely the same thing, albeit almost always fairly modest in size. This evolutionary reduction of the CF muscle along the mammal (synapsid) lineage hasn’t gotten nearly as much attention as that given to the dinosaur/bird lineage’s CFL. Somebody should give it a thoroughly modern phylogenetic what-for! Science the shit outta that caudofemoralis…

Yet, oddly, to give one apparent counter-example, cats (felids) have, probably secondarily, beefed up their CF muscle a bit:

Cats have a pretty large CF muscle in general, and this jaguar is no exception! But mammals still tend to have fairly wimpy tails and thus CF muscles, or they even lose them (e.g. us?).

Cats have a pretty large CF muscle in general, and this jaguar is no exception! But mammals still tend to have fairly wimpy tails and thus CF muscles, or they even lose them (e.g. us?). [photo by Andrew Cuff, I think]

In summary, here’s what happened (click to embeefen):

Better Know A Muscle: The Evolution of M. caudofemoralis (longus)

Better Know A Muscle: the evolution of M. caudofemoralis (longus).

I hope you enjoyed the first BKAM episode!
I am willing to hear requests for future ones… M. pectoralis (major/profundus) is a serious contender.

P.S. It was Freezermas this week! I forgot to mention that. But this post counts as my Freezermas post for 2016; it’s all I can manage. Old Freezermas posts are here.