Luke Horton asked in a comment on a recent post:

Given the chance to examine a titanosaur cadaver with your hypothetical army of anatomists, what would you look for first?

*FACEPALM* How we’ve gone almost 17 years without posting about a hypothetical sauropod dissection is quite beyond my capacity. I am also contractually obligated to remind you that the TV show “Inside Nature’s Giants” shows dissections of a whale, elephant, giraffe, tiger, anaconda, giant squid, etc., so it’s probably the closest we’ll ever get. Go look up photos of Dr. Joy Reidenberg standing, um, amidst a partially-dissected whale, or just watch that episode, and your sauropod-dissection-visualizer will be properly calibrated.

To get back to Luke’s question, there are loads of interesting things that could be dissected in a sauropod, but since the remit here is Matt Wedel x titanosaur, there’s only one possible answer: the lung/air sac system and its diverticula. For several reasons:

Hypothetical reconstruction of the lungs (red) and air sacs (blue, green, and gray) in Haplocanthosaurus CM 879. I’d love to know how close this is to reality. Wedel (2009: fig. 10).

First and most obviously, I’ve spent the last quarter-century trying to infer as much as possible about the respiratory systems of sauropods based on the patterns of pneumaticity in their skeletons, and I’d kill for the opportunity to check the accuracy of my inferences — and those of all my fellow-travelers in the sauropod and dinosaur respiration biz, like Daniela Schwarz and Emma Schachner and Tito Aureliano and many others.

Sauropod respiratory system modeled on that of a bird. I’ll bet the correspondence wasn’t this close. (Also, since making this figure 20 years ago, I’ve learned that the abdominal air sacs of ostriches are actually rather small, although the perirenal, femoral, and subcutaneous diverticula of the abdominal air sacs are extensive; see Bezuidenhout et al. 1999). Wedel and Cifelli (2005: fig. 14).

Second, I am intrigued/haunted by the possibility that extant birds might not represent the apex of saurischian lung/air sac evolution. Birds survived the K-Pg disaster because they were small; respiratory efficiency had little or nothing to do with it (evidence: all the other small-bodied tetrapods that survived, like the many, many squamate and mammalian lineages). To me it would be a wild coincidence if the tiny dinosaurs that survived also just happened to be The Bestest (TM) at some anatomical/physiological thing unrelated to their survival. In fact, given how sensitive birds are to airborne dust and ash, I wonder if their fancy lungs weren’t more of a hindrance than a help in the dusty, sooty, iridium-laced post-impact world. Anyway, there are interesting clues that the air sac systems of extant birds are just one subset of a much greater original diversity, like most (all?) birds starting out embryologically with a dozen or so air sacs, which get simplified to the usual 9 or fewer by fusions. What did other dinosaurs do with their 12 (or more?) air sacs? If any dinosaurian clade was going to push the capabilities of the “avian” lung/air sac system in interesting directions and to fascinating extremes, sauropods seem like a good bet.

Rib articulation angles in the dorsal vertebrae of (a) Lufengosaurus, (b) Diplodocus, (c) Haplocanthosaurus, (d) Tyrannosaurus, and (e) an ostrich. Anterior is to the right. Diplodocus and Haplocanthosaurus are pretty wildly different considering they coexisted in the Morrison. I really gotta write a whole post about that. Boisvert et al. (2024: fig. 12).

So I’m intrigued by the idea that extant birds show us one way that a saurischian lung/air sac system can work, but don’t exhaust the territory, anymore than kangaroos show us all the ways that mammals can reproduce. Maybe sauropods had even better lungs than birds! Or maybe not. Likely they were doing their own weirdly specialized thing — or many weirdly specialized things — that left few to no diagnostic traces in their skeletons. We can be pretty confident that at least some of the pneumatic diverticula of sauropods worked essentially identically to how they do in birds (see Woodruff et al. 2022 and this post), and mid-dorsal pneumatic hiatuses in juvenile sauropods — predicted by me in 2003, found by Melstrom et al. (2016) and Hanik et al. (2017) — suggest that their air sac systems were broadly comparable. On the other hand, the variety of rib articulation angles just within Morrison sauropods tells us they weren’t all ventilating their air sacs in quite the same way (Boisvert et al. 2024), despite broad similarities with other dinos at the levels of rib osteology (Wang et al. 2023) and whole-thorax construction (Schachner et al. 2009, 2011). (Aside: why the hell didn’t I work a citation of Wang et al. 2023 into the Dry Mesa Haplo paper? I can only conclude that I am at least occasionally an idiot.) Whatever was going on, I’m pretty sure sauropods didn’t look exactly like 60-ton turkeys on the inside, but we don’t have a ton of real data on how they differed. It would be amazing to find out.

The mounted Rapetosaurus skeleton at the Field Museum, traced from a photo. Specific weird things to note: neck about twice as long as tail, cervical vertebrae about twice as tall as dorsals, and smallish pelvic bones relative to hindlimbs (= skinny posterior abdomen, at least dorsoventrally). See this post for details.

Third, if any sauropods were going to rival or exceed birds in fancy under-the-hood anatomical and physiological adaptations, my money would be on titanosaurs. They were morphologically disparate, phylogenetically diverse, geographically widespread, they independently evolved to giant size more times than any other sauropod clade, and their growth rates were wild. I’d dissect any sauropod I got access to (uh duh), but a titanosaur would be particularly appealing. Which titanosaur? Probably Rapetosaurus: we know it grew very fast early on (Curry Rogers et al. 2016, and see implications for the nervous system in Smith et al. 2022), it had a highly pneumatic vertebral column (O’Connor 2006), its body proportions were pretty wacky, and it had other features of interest to me, like expanded neurocentral joints (see Wedel and Atterholt 2023 and this post) and neural canal ridges (see Atterholt et al. 2024 and this post).

I used this photo of a Rapetosaurus caudal vertebra a few posts ago to illustrate the neural canal ridges, but — like many other sauropods — it also has very expanded neurocentral joints forming boutons. From Curry Rogers (2009: fig. 27).

Oh, and if I got to dissect more than one sauropod, the rest of my top 5 choices in order would be:

  • the owner of BYU 9024 (Supersaurus? Giant ancient individual of Barosaurus? Are those even different things? Dissecting this critter could tell us!), Barosaurus being the most diplodocid-y and least titanosaur-y neosauropod I know of, and BYU 9024 being from a hellaciously big individual no matter what its classification;
  • the Snowmass Haplocanthosaurus, because I have just so many questions about all the weird stuff going on with its tail (see Wedel et al. 2021 and this post for starters); 
  • Omeisaurus or Xinjiangtitan, to represent a maximally derived-but-also-weird non-neosauropod;
  • Sauroposeidon, for obvious emotional reasons (but not enough to dethrone the others).

After that? Probably Isanosaurus or Melanorosaurus or something else waaaay down the tree, so I could see how much of the sauropod kit was in place from the get-go (probably most of it).

Bone vs joint space in the proximal caudals of the Snowmass Haplocanthosaurus. I’d give one non-essential organ to dissect that tail!

And after the respiratory system, next up for me would be the spinal cord and any related morphological specializations of the neural canal — see Table 3 in Atterholt et al. (2024) for a running tally, and this page. Then intervertebral joints, digestive tract, and reproductive system (neither of the last two leave anything useful in the way of skeletal traces), in that order. Arguably the intervertebral joints would be a bigger score for sauropod paleobiology than spinal cord stuff, but maybe not, and having squelched my emotional pick among sauropod taxa, I’m letting my emotions rule when choosing body systems to dissect. I also am intensely interested in the possibility of protofeathers in sauropods, but you don’t have to dissect those, you can just see if any are present, so I’d cheat a little and note any integumentary specializations en passant. (Remember than an animal can have hairs without being hairy [naked mole rats, rhinos, manatees, dolphins], ditto for feathers.)

So that’s the sauropod and the body system I’d dissect first, if given the chance. What’s your answer?

References

 


doi:10.59350/ajsh7-42642

It is a measure of how scattershot our blogging is that we haven’t mentioned Adam Mastroianni or his blog Experimental History before now. Mike and I enjoy a lot of freedom from institutional oversight of our research and publishing, and we do make some use of it. We like to try new publication outlets — we published the first paleo papers in Qeios, Academia Letters, and, most notably and satisfyingly, PeerJ — and we occasionally like to imagine a world of scientific inquiry that was less encumbered by the cruft of bureaucracy, tradition, peer review, the perversion of impact factors, and the deliberate roadblocks thrown up by barrier-based publishers.

Mike and I both admire Adam Mastroianni because he just went and freakin’ did it. Quit his academic job to live off his Substack subscriptions, published a paper by posting it freely to the web (along with the supporting data [and yes, I know he’s not the first to do this; it’s still cool]), and not only envisioned a new way of doing science (Science House!), but actually implemented it.

Oh, and we also admire him because he thinks interesting thoughts and his writing style is right up our alley — light-but-direct, conversational, self-deprecating, insightful, and often wickedly funny. He makes it look easy, which is virtually always the visible result of a lot of hard work. (Unfortunately, it is also possible to work hard without making anything look easy.)

Lest you think I’m ladling on the compliments because of what’s coming up, I’ve probably made a nuisance of myself among my close friends and colleagues by forwarding Adam’s posts a LOT this past year. I’m now having conversations with friends of the form, “Have I sent you anything by Adam Mastroia-” “YES.”

ANYWAY, earlier this summer Adam announced a blog-writing contest, for the lovely purpose of helping him find new reading material. That announcement led to the following exchange.

Matt: Thinking this might be the prod I need to finish “We’re not going to run out of new anatomy anytime soon”.

Mike: That’s a good call. Maybe a bit TOO obviously Mastroianni-bait?

Matt (somewhat later): Apparently just the right amount of Mastroianni-bait — he just wrote to tell me that post won the contest!

I did make one tiny change from the version submitted on Google Docs and the version posted here — I added a sentence about the new neural canal ridges paper (Atterholt et al. 2024, this post, and this one), a paper which nicely illustrates the thesis of the new anatomy post. It’s all come together very satisfactorily: at SVPCA in 2019, Jessie gave a talk on neural canal ridges, and my talk, which I think came immediately after hers, was about how to make new discoveries in human anatomy. Here we are five years later to the month, and the neural canal ridges paper and the long-in-development blog post about new anatomy are both finally out within a fortnight of each other.

Congratulations to the other winners! I’m looking forward to digging into the spring tide of interesting reading that Adam’s contest aggregated, listening to new voices, and having new thoughts. The excerpts and descriptions in Adam’s reveal post are beyond intriguing — go check ’em out.

And a huge thank you to Adam — for writing Experimental History in the first place, for showing the rest of us that there are alternatives to the Way Things Have Always Been Done (hint: “Always” in that formulation is a much shorter period of time than most people think), for running the contest and introducing so many interesting writers to each other and the world, and, of course, for choosing my entry. The world gives me a lot of positive feedback already, but mostly for the “official” stuff like publishing and teaching. It’s really nice to get recognized for blogging, especially out of such a rich field of entrants.

If anyone’s curious, I’m going to use the prize money to fund my research. I drew out my startup money for a loooong time but it’s finally gone, I don’t have a current grant and applying for grants is a not-fun way to gamble weeks or months I could use to write papers, and the little pot of annual travel and research money from my department only goes so far. The prize money will let me keep finding and reporting on new anatomy. Probably mostly in sauropods, but who knows — I’m publishing (and blogging!) on freakin’ fish these days, so anything’s possible. Stay tuned! (And welcome, Experimental History readers!)

All right, that’s plenty of navel blogging. Back to the good stuff in the next couple of posts.

Reference

 


doi:10.59350/vha9z-9km17

[This post received first place in the 2024 Blog Extravaganza at Adam Mastroianni’s Experimental History. Many thanks, Adam!]

I first had this thought in 2019, and I started this draft in early 2020, but…you know how that particular story turned out.

I’m picking it back up again now because I’ve had the titular point reinforced on several trips and projects over the past couple of years. And because I think it’s ultimately a hopeful message. If you are interested in making anatomical discoveries, good! Because relative to a single human life, the work to be done is effectively infinite.

But wait, you might say, how could that possibly be true? Have we not been plumbing the depths of the human body literally for thousands of years? Have we not imaged people down to micron resolution with every available scanning modality?

We’ve been at this a while, how are we not done yet? Left: Da Vinci. Right: Hua Shou, 1340s, Ming Dynasty.

And what about other extant critters? Chickens are one of the commonly-used model organisms in laboratory studies, and the basis for a multi-billion-dollar food industry. Surely we must know everything there is to know about their anatomy? (Spoiler alert: we do not.)

What about fossils? Are we not even now engaged in a massive, civilization-wide, distributed project to scan museum collections? Can we not publish entire dinosaur skeletons as 3D files in the supplementary information to our papers (Lacovara et al. 2014)? There will always be new fossils to discover, but can’t we at least say that the ones we’ve digitized are completely known?

Where is all this new anatomy hiding?

I’ll tell you.

(Warning: dissection images inbound. Nothing too gory, but still.)

I’m going to draw a lot from human anatomy, because it’s one of the areas where I have the most hands-on experience, and because humans are one of the best-studied organisms on the planet. So if there are macroscopic structures awaiting discovery in humans, imagine how much more true that will be of every other species that we haven’t been studying with extreme diligence and self-interest for millennia.

The Human Factors

Part of the reason why we are still making new discoveries in human anatomy is because we’ve made the process of finding, recognizing, and publishing new structures surprisingly difficult. None of these barriers were put in place deliberately (we could quibble about barriers to publication), but they’re slowing the advance of anatomical knowledge nonetheless.

1. Not everyone gets to look, and everyone who does is on the clock

I’d originally put this point farther down, but for human anatomy it is the subtext and background radiation for everything else I have to say, so I’m giving it pride of place.

When we described the long cutaneous branch of the obturator nerve a few years ago (Staples et al. 2019, this post), I wondered why it hadn’t been discovered sooner. I hypothesized that it fell into a perceptual blind spot: the people with the best chance to discover it were medical students and surgeons, and each group faced a significant barrier. Surgeons had the expertise to recognize and preserve this tiny, delicate nerve, but they didn’t have the time or operative freedom to flay their patients open from ankle to groin to trace its path. Med students had the opportunity to chase the variant nerve all the way down the lower extremity, but only if they managed to preserve it while skinning the limbs, and if they recognized it as anomalous – neither of which was likely on Day 1 when they did the skinning.

Preserving that very long, very skinny nerve in dissection is not easy. Modified from Staples et al. (2019: figure 5).

Later I realized that these same factors apply to all kinds of anatomical discoveries. No shadowy Illuminati group deliberately made this decision, but as a civilization we have collectively ‘decided’ that three groups of people would get to peer inside the human body, and they’d all be hobbled. Surgeons are under immense pressure to make smaller incisions, do less invasive surgeries, and keep their patients on the table for as little time as possible, because small holes and short surgeries generally correlate with better outcomes. I’m not saying this is wrong – it is undoubtedly the right decision in the vast majority of cases – but it does mean that our most experienced anatomists have very little opportunity to investigate possibly new anatomical features, unless they happen to impede a surgery.

The second group that gets the privilege of hands-on exploration of the human body is medical students, and they’re also on the clock. Med school is legitimately challenging – we use the metaphor “drinking from the firehose” a lot – and med students usually have a long list of structures to find in a 3-4 hour dissection. I don’t think anyone could reasonably blame med students for not being “discovery oriented”; the fact is that when you’re going to spend between 100 and 200 hours dissecting an entire human body, at some point it becomes a job, and with all the other subjects med students are expected to master (biochemistry, cell biology, physiology, microbiology, pharmacology, etc.), it’s not their only job, and not always the top priority in a given day or week.

That leaves the third group: anatomy teachers, like me. With dozens or hundreds of med students to do the dissecting for us, shouldn’t we be in a perfect position to recognize interesting things in the anatomy lab? To some extent, yes, but the clue is in the question. I’m in the anatomy lab to teach, and teaching a big room full of very smart, very motivated folks who have Wikipedia and Radiopaedia and their textbooks and the campus library on their phones and tablets is a bit of a high-wire act, requiring dedication and focus – on teaching, not on discovery. So I keep my antennae out, probably more than most, but I’m still relying on the med students to make the discoveries, and I suspect that is true of most anatomy teachers.

2. Anatomical knowledge is oddly siloed

If you crack open the 40th edition of Gray’s Anatomy, published in 2008, and turn to page 1419, the very first sentence about the fibularis tertius muscle reads, “Fibularis tertius (peroneus tertius) is a muscle unique to humans.”

That bold assertion would probably come as a surprise to Dudley Morton, who published a paper titled “The peroneus tertius muscle in gorillas” in The Anatomical Record…in 1924. And to William Straus, Jr, who described and illustrated the peroneus tertius muscle in chimps and gorillas in a 1930 paper in The Quarterly Review of Biology.

Morton (1924), first page and figure 1.

How did this happen? The Anatomical Record and The Quarterly Review of Biology are not obscure sources, they’re highly-regarded journals with global readership. There’s a long story here, involving the prominent (not to say tyrannical) Victorian anatomist Richard Owen, the Gilded Age quest to tally anatomical features separating humans from apes, and some extremely dubious evolutionary hypotheses, but the short version is that comparative anatomy, physical anthropology, and clinical anatomy are three distinct fields. Each field has its own preferred publication venues, citation classics, and bodies of “common knowledge”. Ideas that were sunk long ago in one field may still be viable in another, because the debunking happened in a paper that few people outside of its home field have ever read or cited. And not just hypotheses, but even basic facts, like whether the peroneus tertius muscle is actually unique to humans (for avoidance of doubt, it most certainly is not).

This weird balkanization of science doesn’t make it harder to spot anomalous and potentially new anatomical structures in the dissection lab, but it can impair people’s efforts to understand the evolutionary history and clinical importance of a given body part, especially if they happen to fall into one literature silo and never learn that the other, parallel ones exist.

3. There are many barriers to publication

Crucially, both surgeons and med students live notoriously busy lives. Even if they notice and preserve something interesting, plowing through the literature, getting publication-quality photos, and actually writing and formatting a paper all take time. Hardly anyone has the time to do all the work by themselves, but collaboration means coordinating the efforts of multiple busy people. Then there’s the task of finding an appropriate journal – loads of otherwise promising OA outlets don’t take anatomical case studies, for example. And finally there is the gauntlet of peer review, about which we’ve already spilled many words.

A slide from my 2019 SVPCA talk, “How to make new discoveries in (human) anatomy.”

Now, in point of fact, surgeons, med students, and anatomy professors do find and publish new anatomical discoveries. But there are enough hurdles just on our side to explain why we’re not done yet, and may never be.

Nature Doesn’t Make It Easy

Beyond the speed bumps we humans have accidentally erected there lurks the unending, phenomenal complexity of nature, which throws up its own barriers to discovery.

1. Humans and other animals are hideously complex

Dissection-based human anatomy courses run between 100 and 200 hours not because that’s an administratively or pedagogically convenient number – I and everyone else in medical education, and especially the bean counters, can assure you it is not – but because that’s simply how long it takes to find all the bits. Minimally – we expect that students will take advantage of open lab hours on evenings and weekends to tidy up their dissections. And that’s relatively hasty, on-the-clock dissecting for teaching purposes. The professionally prepared plastinated cadavers for exhibits like Body Worlds can require 500 to 1000 hours of dissection.

That might sound ridiculous. After all, professional butchers, and hunters and farmers who dress their own kills and livestock, all get very good at taking apart large mammals much faster than that. But there’s a world of difference between taking apart a carcass as efficiently as possible – for which I give all those folks full props – and trying to dissect and put a name to all the parts.

Esophageal plexus and other neck viscera in left lateral view. For more about that variant nerve, see this post. Altounian et al. (2015: figure 4).

I was confronted with the frankly appalling complexity of the human body about a decade ago, when as part of a student project (Altounian et al. 2015) I did a deep dissection of the esophageal plexus. I went in after hours to do the extra dissecting work, just like we encourage the med students to do, and it took me something like four hours. It was rewarding work, but it’s probably telling that in ten years I’ve never done it again.

Incidentally, I don’t think this gets much easier as animals get smaller. A chicken or a cat has about the same number of body parts as a human, they’re just smaller and harder to see. Frogs seem to be a little simpler than shrews or hummingbirds, but it may also be that we know them less well, and dissect them less patiently and completely. At some point gross anatomy has to give way to histology as body parts become microscopic, but that doesn’t mean that the animals in question aren’t still pretty darned complex.

In sum, humans and other animals have lots and lots of parts. But it gets worse.

2. Anatomical variation is extremely common

It took me a long time to realize that there’s a needle-and-haystack problem with recognizing genuinely new anatomical structures from the common variations that turn up all the time. This is one of those things that might seem hard to believe unless you’ve experienced it, but we humans are crazy variable under the hood. In my program we encourage the students to log interesting variations on the whiteboard in the lab, not least so that everyone can beware of the variant anatomy while studying for their practical exams. If the students are really diligent about the logging, something like a third of the donor bodies end up written on the board. And those are the variations the students found and worried might distract their studying, not all the variations that exist. Oh, and we reset the log between each of our five curriculum blocks through the year. So essentially every cadaver has a chance to end up on the ‘variation board’ at least once, and some might be up there three or four times.

Here’s why this is relevant: numerous times I’ve seen some variation in lab, in a body system or region in which I was not familiar with the primary literature, and I’ve thought “cool variation” and moved on. Then later I’ll get curious and look it up, or I’ll be researching something completely different and stumble over a mention of that same variation. A couple of times that variation has turned out to be so phenomenally rare that if I’d only gotten good photos at the time, I could have had a nice little paper.

So to a first approximation, almost every human being has at least one anatomical variation notable enough that a med student would write it on a whiteboard. And this is actually a problem, because those of us who work in anatomy labs see so many of those common variations that sometimes it keeps us from recognizing the truly novel and important stuff.

3. Some body parts have distractors

What we now call the anterolateral ligament (ALL) of the knee was first discovered by a French surgeon 145 years ago (Segond 1879), and independently rediscovered sporadically throughout the twentieth century, but it wasn’t widely recognized as a body part normally present in most people until a pair of papers in 2012 and 2013 brought it to global prominence (Vincent et al. 2012, Claes et al. 2013).

A diagram from my 2019 SVPCA talk, showing the ALL (red) sandwiched between the patellar retinaculum and the iliotibial tract.

Given the vast amount of time, money, and effort that humankind has put into understanding, rehabbing, rebuilding, and replacing our knee joints, the absurdly long period during which the ALL escaped wide detection is flat-out amazing to me. But it also makes sense in a weird way. The ALL angles downward and forward from the lateral aspect of the distal femur to the anterior portion of the proximal tibia (hence anterolateral ligament), and in that position it is sandwiched between the patellar retinaculum, which lies deep to the ALL, and the iliotibial tract, which lies on top of it. Crucially, both the patellar retinaculum and the iliotibial tract are made of dense connective tissue, like the ALL, and they run in the same direction as the ALL.

I’ll bet that in the decades and centuries before the 2010s, hundreds if not thousands of surgeons and medical students saw the ALL and mistook it for part of either the patellar retinaculum or the iliotibial tract – structures that they were expecting to see in that region, also made of connective tissue, also running in the same direction.

If you only get to look inside the box, these two things appear identical. Modified from Staples et al. (2019: figure 6).

A similar thing probably happens with the aforementioned long cutaneous branch of the obturator nerve. In the two known cases, it was running with the great saphenous vein, in a position usually occupied by a branch of the saphenous nerve. I reckon that surgeons see the long cutaneous branch of the obturator nerve on a regular basis, but they have no way of knowing that it’s a weird variant because it sits where they were already expecting to see a nerve.

It’s hard to say how important this factor is, but I note that almost all the anatomical variants I’ve helped students present at conferences or publish are things that they found in complicated areas – nerve plexuses, bundles of tendons crossing a joint, and so on – where they could easily have escaped detection if people hadn’t really been on the ball. And of course I can only count the hits; I can’t tally all the variants that we missed because we mistook them for their distractors. Thoughts like that haunt me.

4. Some things are just hard to see

The plain fact is that some parts of the body are easier to investigate than others. I’ve written a lot about how the pneumatic diverticula of birds are under-documented, even in chickens, the most numerous and best-studied birds on the planet (whinge, whinge). But diverticula can be surprisingly tricky – when birds die, many of the diverticula empty out and collapse. The diverticula can look just like loose connective tissue, unless they’ve been injected with latex or resin, or re-inflated and CT scanned, and both the injection and the scanning take a lot more time and effort than a simple dissection. One handy thing about the paramedullary (or supramedullary) diverticula is that they’re unable to collapse; the bony walls of the neural canal keep them propped open whether they’re inflated or not.

An ostrich neck in cross-section, showing many of the pneumatic diverticula of the respiratory system. The neural canal is the bony tube around the spinal cord. From this post.

Speaking of, the neural canals of archosaurs host a whole zoo of anatomical novelties – big veins, pneumatic diverticula, odd joints, ligament scars, and, oh yeah, an entire novel balance organ. Although the big veins (in crocs and some birds) and pneumatic diverticula (in many birds) have been known to exist since the 1800s, they’ve really only started to be adequately documented in this century. The same goes for everything else on the list; to pick a timely example, the ligament scars were only described for the first time in archosaurs a couple of weeks ago. Why the delay? I think that neural canals, being relatively small-diameter bony tubes, are just that much harder to study than most other parts of the body, whether we’re talking about big-ass crocs or tiny hummingbirds. Heck, one of the most recently-discovered macro structures in the human body is the midline interlaminar ligament, only recognized for the first time in 2019 (Simonds et al.), which lies – you guessed it – along the roof of the neural canal.

So one way to make new discoveries is to simply look in inconvenient places. Sacral pneumaticity in dinosaurs is poorly understood because the sacral vertebrae are often inaccessible, but there are ways around that: studying the unfused sacral vertebrae of juvenile and subadult animals, looking at broken specimens, and staying alert for interesting opportunities. But now I’m getting ahead of myself – problem solving deserves a whole section.

What to do about it

Of the factors slowing down the pace of anatomical discovery that I numbered above, all but the first can be overcome with time, curiosity, patience, and determination. One of the biggest boosts is simply being aware that new discoveries are still being made, and staying on the lookout for them.

As for the first – the fact that not many folks get to dissect human bodies, and everyone who does is busy – I could fix that if I was sufficiently rich. If I was a multi-billionaire, I’d hire 1000 of the world’s best surgeons (in staggered waves, so I didn’t doom thousands of patients by pulling too many experts off the line at once), supply them with 10,000 ethically donated willed bodies representing as many geographic regions and genetic backgrounds of humankind as possible, and give each surgeon a couple of years to dissect their 10 bodies, ideally in labs with 50-100 bodies at a time so the small groups of surgeons could look at each other’s work without getting overwhelmed, or work in teams if they preferred. I’d also supply them with professional photographers to document everything they found, and a small army of research assistants to help them with library work and writing up. That wouldn’t be enough to declare the science of human anatomy a completed project, but we’d know a heck of a lot more than we do now.

I’m not a multi-billionaire, and no-one on the planet is ever going to fund the vast study I just described. I think we’ll still get to an equivalent level of knowledge, but it will take the next 500 to 1000 years, as those discoveries are made piecemeal, mostly by alert medical students who happen to do better than average dissections in their gross anatomy courses.

Turning to comparative anatomy, I’ll conclude this section with one of my favorite published sources. Baumel (1988) is a 123-page book on the anatomy of the tail of the pigeon. If a chunk of pigeon the size of the last digit of one of your fingers can bear over 100 pages of detailed examination – and it can, I have the book and I refer to it in my research – then we are not going to run out of new anatomy anytime soon (not least because there are the other 10,000+ species of birds that have not had their tails described in that level of detail).

But is it worth it?

Sure, people might say, some goobers might write boring-ass treatises about pigeon tails or chameleon tongues or frog pelvises, but isn’t that all just so much pointless stamp collecting? Does any of it really matter? Shouldn’t we funnel our limited support for science toward things that are going to make a practical difference?

I’d counter that science is a young enterprise and we are still exceptionally bad at determining in advance what kinds of things are going to be important in the future. Baumel’s book on pigeon tails has been cited just in this decade in fields as diverse as biomechanics, embryology, evolution – and, hey, by researchers investigating rudderless flight control for UAVs, a technology application that didn’t exist when the book was first published. The skin of sharks inspired wetsuits so efficient they’ve been banned at the Olympics, and the first-in-class COVID-19 medication remdesivir is one of hundreds of pharmaceuticals derived from the biochemistry of sea sponges. I think documenting the universe is a noble goal in itself, but we should probably keep researching All the Things because that’s where the new technology is going to come from. And the people – nations, states, businesses, inventors – smart enough to invest in basic science are going to get those discoveries before anyone else does.

And anyway, compared to most other fields of inquiry, anatomy research is dirt cheap. Embalmed human cadavers cost money, but I could still get the 10,000 cadavers I’d need for my dream project for less than the cost of a Marvel movie. Of course that project is never going to happen, but fortunately we can continue piggybacking human anatomical research on the vast anatomy education effort necessary to train physicians. For comparative anatomy and paleontology, we basically need to keep giving geeks a little research time and a ten-thousandth of a percent of the cost of the Large Hadron Collider so they can keep themselves busy when they’re not teaching or running museums, and they’ll keep doing the work. (That’s not to say that more support wouldn’t be appreciated, or speed things up a little.)

So if you like anatomy, come join the hunt. You probably won’t get rich, but you’ll stay busy doing interesting work, which is a different form of wealth. And if you stay alert, you will not run out of new things to find.

References

 


doi:10.59350/63r4z-32f49

Trunk vertebra of a tuna (Thunnus), OMNH RE 0042, showing paired bony spinal cord supports

Here’s a grab-bag of follow-up stuff related to our new paper on neural canal ridges in dinos (Atterholt et al. 2024, see the previous post and sidebar page).

Neural canal ridges, or bony spinal cord supports?

I got into the habit of calling the inwardly-projecting bony prominences in the neural canals of sauropods and other critters “neural canal ridges” partly because I was thinking about them for literally years before I knew what they were, and I had to call them something, and partly because “neural canal ridges” is a reasonably accurate descriptive term that does not imply a specific function. NCRs became part of my internal lexicon.

Later on, thanks first to David Wake, and later to Skutschas & Baleeva (2012), we discovered that extant fishes and salamanders have bony spinal cord supports, and we think that’s the best explanation for why NCRs show up in so many dinos. “Bony spinal cord supports” is not function-neutral, it takes a stand. Since the whole point of our paper is not only to describe these things in dry terms, but to also take a stand on their associated soft tissues, it would be more coherent to cowboy up and call them “bony spinal cord supports” instead of “neural canal ridges”, and that’s exactly what Jessie Atterholt did in the tables and figure captions of the new paper. Also, sometimes the bony spinal cord supports are not ridges, but shelves or planks or spikes — check out that tuna vertebra up top, and the salamander verts in Fig. 1 of the new paper — so “neural canal ridges” doesn’t even accurately describe them all the time. If I call them NCRs in my blogging, it’s out of habit, and because — so far — that does accurately describe the appearance of the bony spinal cord supports in dinos.

Denticulate ligaments: sometimes double, sometimes absent

Here’s something that turned up late in our research on this project. Elvan et al. (2020) is a nice paper on the denticulate ligaments in developing humans (it is of course tragic when fetuses are miscarried or stillborn, but what we learn from them can help keep others alive). One of the curious things they mention, and figure, is that the denticulate ligaments that suspend the spinal cord inside the dura mater are occasionally doubled on one side, and occasionally absent.

Elvan et al. (2020: fig. 1)

This shouldn’t be super surprising. Variation exists in part because developmental programs are messy. “Asymptomatic anatomical variation”, “pathological variation”, “congenital anomaly” (“birth defect”), and “fatal malformation” are points on a spectrum — and all of us are somewhere on that spectrum. “Normal” human anatomy is normal in the statistical sense, in that the majority of folks end up in the big middle, but that middle encompasses a lot of variation, and there are long tails in lots of directions for almost every body part and body system, and things can sometimes be pretty non-standard under the hood without causing noticeable symptoms.

Here’s a whole paper on a six-legged rat (Brown 1996). Click to embiggen.

In particular, if there’s a developmental program for building structure X — whether structure X is a hair follicle, a muscle, nerve, or blood vessel, a finger or toe, a gill arch, a vertebra and its associated body segment, or an entire limb — then inevitably there will be counting errors from time to time, omissions or duplications, and embryos, fetuses, or offspring produced with fewer or more of structure X than is typical. At the small end of the scale we might not even notice, and at the large end of the scale the variation might not be viable.

In between those extremes you sometimes get a memorable villain.

ANYWAY, finding the Elvan et al. paper was an “Aha!” moment for me. Back in 2018 when I’d been photographing tuna vertebrae in the OMNH collections, I found some that had not one but two inward-pointing bony spikes on each side. I figured these were just a fancier system of bony spinal cord supports, probably indicating doubled denticulate ligaments. I didn’t know for sure that the latter existed, so in assembling figures for the paper we went with the tuna vertebra that most closely resembled the salmon vertebra figured by Skutschas & Baleeva (2012). Later on, the Elvan et al. paper confirmed for us that doubled denticulate ligaments sometimes occur, at least in humans, so it’s plausible that they happen in fish, too, and maybe regularly given that I found the quad-spike setup in multiple tuna vertebrae. But that seemed like a lot of extra yap and figures to make a rather minor point, which is why you’re hearing about this in a blog post instead of in the paper.

Another vertebra of OMNH RE 0042, showing (what I infer to be) paired bony spinal cord supports

I assume that these spikes and whatever attaches to them were described back in the 1800s in some obscure paper, probably published in Germany or Great Britain, but if so I’ve not yet tracked down that hypothetical publication. Even if said publication exists, I’m sure it’s illustrated with a hand-drawn diagram. It occurs to me that someone could go to a fish market, buy a chunk of tuna with the bone in, do a little careful dissecting, get some hi-res color photos, and have everything they’d need to publish a nice little paper, either describing these spikes and their soft-tissue correlates for the first time, or redescribing them and providing the first good color photos. Realistically I’m unlikely to get around to that, so if you want it, go nuts.

Science…and dinner

Citing the Deep Magic

I’m gonna geek out for a sec on the developmental underpinnings of the denticulate ligaments and the vertebrae they’re associated with. And to do that, we have to orient ourselves to the various bits sticking out of the spinal cord and how they relate to the vertebral column.

Here’s a chunk of sauropod tail in left lateral view (modified from Wedel et al. 2021: fig. 2a) — specifically, a 3D-printed section of Haplocanthosaurus tail that Alton Dooley put together for the “Tiny Titan” exhibit at the Western Science Center a few years ago, seen in medial view in the second image down in this post. The laterally-facing bony loop formed by the central and zygapophyseal articulations of two adjacent vertebrae is the intervertebral foramen, and it’s through the intervertebral foramina that the spinal nerves leave the neural canal (blood vessels enter and leave through these openings, too). Assuming that sauropods were built like reptiles rather than mammals, and lacked epidural fat, a horizontal section through this bit of tail on the black line indicated by the Xs might look something like this:

Anterior is toward the top now. There’s a lot going on in this image, so let’s take it one piece at a time. The neural arch pedicles are the paired black-and-white pillars on either side of the spinal cord, defining the lateral walls of the neural canal. (The section in the photo also went through the caudal ribs but I was too lazy to draw those.) The meninges — the dura, arachnoid, and pia mater, and the subarachnoid space — are by now old friends; this diagram is showing us the same structures as this one from the previous post, just in horizontal section rather than transverse. Bundles of spinal nerve roots come together to form the spinal nerves, which exit the neural canal at the intervertebral foramina between adjacent neural arch pedicles. The various meninges form little sideways-projecting meningeal sleeves over the first little section of each spinal nerve; imagine making 3-layer coveralls for a centipede and you’ll have a good mental model of the whole meningeal system of the spinal cord (for real geekery, past the ends of the meningeal sleeves the nerves are jacketed in a different connective tissue called epineureum). The denticulate ligaments attach the spinal cord to the dura mater (or even through the dura mater) level with the neural arch pedicles of the vertebrae, so if you’re looking at a section of the cord in dorsal or ventral view you’ll see bundles of spinal nerve roots (at the intervertebral foramina) alternating cranio-caudally with denticulate ligaments (in between intervertebral foramina). You can check that with the dorsal-view photos of human spinal cords above and in this image in the previous post.

(Note for any confused med students who might be reading this: anatomical position for humans is upright, so horizontal and transverse sections are synonymous. Most other animals carry their bodies horizontally, so a horizontal section through a sauropod would be similar to a coronal or frontal section through a human vertebral column. Also, humans do have epidural fat, unlike this sauropod, and our denticulate ligaments do not go through the dura mater to attach to bone. So don’t use these sauropod diagrams to study for your human anatomy courses! Instead, a great learning exercise would be to redraw this diagram so it was accurate for a human. If you do that, feel free to drop me a line in the comments and we can talk about your results. Standing offer, good forever.)

At the bottom of the image I labeled segmental muscles and intermuscular septum. You’ve seen these before, although you may not have known it: they make the zig-zag patterns in the meat of fishes, where we call the segmental muscles myomeres (“muscle parts”) and each intermuscular septum a myoseptum, plural myosepta (“muscle partition”).

Lateral view of the trunk muscles of a salmon, Salmo. Liem et al. (2001: fig. 10-16)

Each myomere is associated with a particular spinal level — a paired set of spinal nerves, like the C7 or T10 spinal nerves in a human — and each myoseptum is associated with a particular vertebra, like, er, C7 or T10 in a human (or a sauropod, although we’d call it D10 for dorsal 10 in a sauropod; sauropod dorsals all have big ribs that were mobile at some point, so there’s no need to separate them into thoracic [dorsals with mobile ribs] and lumbar [dorsals without mobile ribs]). Put a pin in that thought for a moment, we need to wrap up something fishy.

Myomere cones in a salmonid, Salmo (A), and a dogfish, Squalus (B, C). Liem et al. (2001: fig. 11-4).

You maybe looking at the mild zig-zaggy-ness of the myomeres in that first salmon diagram, and the target-like concentric circles in the photo of the salmon steaks up above, and thinking something doesn’t add up. And you’re right — the surface zig-zaggy-ness of the myomeres is not their full extent, they have anterior and posterior cones arranged concentrically, presumably to allow each myomere to exert force over more of the vertebral column. And that’s why fish comes apart in such interesting ways when you eat it, especially if it’s cooked.

Anyway, back to the segmental muscles and intermuscular septa in the sauropod — and in yourself, for that matter. It’s not immediately obvious that amniotes are built on the same myomere/myoseptum infrastructure as sharks and salmon, because our development involves a lot of splitting and recombining and stretching of muscles across multiple spinal levels. But if you go deep enough, we all have some single-segment muscles that bridge adjacent body segments — intercostal muscles between our ribs, and interspinales, intertransversarii, and rotatores breves between adjacent vertebrae.

The relevant slide from my lecture on deep back muscles. Rotatores aren’t shown because I’d covered them on a different slide, with the rest of the transversospinal group. I should do a whole post on them sometime.

Now here’s the part that I think is awesome, what this whole section has been building toward: the myomeres and myosepta were there from very early on in development, and the myosepta originally ran from spinal cord to skin. Denticulate ligaments are just what we call the little stretch of myoseptum between the spinal cord and the dura mater, sorta like how we use ‘Foothill Boulevard’ for the stretch of US Route 66 that runs through Claremont and adjacent townships. The pedicles of the neural arches — in fact, the entire left and right halves of each neural arch — form within the myosepta. The light gray boxes around “denticulate ligament”, “neural arch pedicle”, and “intermuscular septum” in my cross-sectional diagram above unite the different portions or aspects of the embryonic myoseptum. I didn’t work all this out myself, mind, I learned it from Skutschas & Baleeva (2012), who demonstrate it all very convincingly with developmental work on larval salamanders.

And that brings us to the weirdness of mammals.

NCRs? No thanks, we’re mammals

I’ve gotten some questions about whether mammals could have NCRs. I doubt it. Not to put too fine a point on it, but as a species we just care more about our own anatomy and that of dogs and cattle and rabbits and rats, than we do about any other critters, and I think if mammals had NCRs they’d have been found and logged by now.

Also, I don’t think we mammals have the capacity to have bony spinal cord supports, because those are the attachment scars of the denticulate ligaments to the inner walls of the neural canals, and our denticulate ligaments don’t work that way. Our denticulate ligaments connect our spinal cords to our dural sacs, but we have epidural fat between the dura and the neural arch pedicles, and apparently when in development the dura pulls away from the neural arch pedicles and epidural fat starts to be laid down in between, whatever embryonic connection existed between the denticulate ligament and the rest of the myoseptum is broken.

I said “I doubt it” rather than a flat “no” because apparently there is very little to no epidural space in the cervical region of most mammals. IF there are mammals in which the dura mater fuses to the periosteum in the cervical region, then maybe the embryonic myoseptal connection could be maintained, the resulting denticulate ligaments could be tied down to bone, and bony spinal cord supports could exist. I wouldn’t rule it out, because if there’s one thing we as a species are even worse about than caring about non-mammals, it’s peering into neural canals.

But we’re working on it.

References

 


doi:10.59350/c8g24-ppe24