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Posts Tagged ‘funky feet’

In case you haven’t heard, Saturday, September 22nd, 2012 (today, at this writing) is World Rhino Day! The main websites include here and here.  Ivan Kwan has also posted a fantastic blog entry “Rhinos are not prehistoric survivors” for WRD2012- check it out! And if you haven’t seen the WitmerLab’s AWESOME Visible Interactive Rhino site, you really really need to (in fact, quit reading this and go there first; it is soooooo good!).

I’ve written about the global rhino crisis before, and about rhino foot pathologies. The title of today’s post may be “cute”, or at least goofy, but the real situation is as grim as the images I’ll share. I won’t repeat the explanation, but all five living species of rhinoceroses are in serious trouble. There’s a good chance that most or all of them will go extinct quite soon– see the previous links for more information on this. Javan and Sumatran rhinos are dangling the most precariously over the precipice of extinction. My goal in this post is to share the beautiful, complex and exotic anatomy of rhinoceros anatomy and movement, and the joy of contributing new scientific information about poorly understood species.

Stomach-Churning Rating: 7/10– dissections, and there are a couple of pics where the specimens are not so fresh, and there’s big skin, and a huge heart.

Baby white rhinoceros. Will frozen specimens like this be all we have of rhinos someday?

The purpose of today’s rhino post is to share a bit more; especially images; of the work my team has done on rhinoceros gait and limb anatomy; all of it unpublished but hopefully coming soon. We’ve steadily been collecting data since ~2005. Because my previous post went through some of this, I’ll keep it brief and image-focused.

First, a video of one of our amusing encounters with a white rhinoceros, at Woburn Safari Park. In this study, we wanted to measure, for the first time really, the gaits (footfall patterns) that a white rhinoceros uses at different speeds, and how often it uses those different gaits. We attached a GPS unit on a horse surcingle around the rhino’s torso, which measured the animal’s speed once a second. We then observed 5 individuals (1 at a time over various days), following them in my station wagon (estate car) across the safari park. We filmed them with a conventional camcorder to document their gaits, and concentrated on the two periods of the day that they’d normally be active: when released from their overnight barn, and when coming in for the night back to that barn. They got rather excited and frisky some of those times. The GPS belt then kept recording speeds for the rest of the day; unsurprisingly, the rhinos generally did not do much. I have to thank Nick Whiting, rhino handler, for his help making this research happen. I’ve been meaning  for too long to finish the final paper… soon, I hope! Enjoy this tense scene of a rhino investigating my car (driven by me and with an undergraduate student filming) then having a nice canter/gallop across the field (accompanied by my jubilant narration).

Like our foot pressure research, we aim that this work provides baseline data useful to caretakers of rhinos; for example, to test if a particular animal is lame. This follows what we’ve successfully done with elephant gaits and feet, translating basic research into more clinical application. But my major scientific interest is in understanding more about what makes any rhinoceros, even a 2-tonne White rhino, so much more athletic than any elephant (even a baby or 2-tonne small adult Asian elephant). As the video shows, they can use a variety of gaits including cantering and galloping, and trotting at slower running speeds. No elephant ever does that, and no one knows precisely why. The leg bones are more robust, but the muscles aren’t that dramatically larger in rhinos.

An Indian rhinoceros forelimb- note the characteristic knobbly hide, unlike the smoother, more elephant-like hide of a White rhinoceros.

Similarly, the anatomical work we do with rhinos is intended to not only be useful science for comparative biologists like me, showing how rhino limbs work and how they differ from those of other animals, but also to aid clinicians in comparing normal vs. pathological anatomy. For conveying that anatomical work, I’m lucky to have been granted permission to use a professional photographer’s pictures of some of my freezers’ rhino specimens– big thanks to James King-Holmes and the Science Photo Library. The watermarked images below belong to them. I ask that you do not use them elsewhere, honouring their license to me for personal usage on this website (and I will only use them here). I’m in all the images, which makes me feel weird putting them up here, but it’s about the rhinos (and freezers), not me. First: the infamous “rhino foot freezer”, featuring some of its denizens:

Second, a re-introduction to multifarious contents of Freezersaurus, but this time featuring rhino feet (here, a skinned white rhino foot that we had already studied):

…and inside we go (and I begin to get frosty and numb-fingered from holding a foot; my smile soon fades):

Taking a rest with the skinned white rhinoceros foot:

And now warming up at the “digital freezer”, our CT scanner, and preparing to scan another rhinoceros foot, which segues nicely out of this image sequence:

Now over to some 3D anatomy– segmented reconstructions of rhinoceros fore (top) and hind (bottom) feet, from CT scans; if you’ve frequented this blog you know the drill. Here, the longest bones are the metacarpals/metatarsals and the upper bones are the carpals/tarsals, then the bones near the botttom are the phalanges, which connect to the hooves (visible in the bottom image):

I’ll wrap up with a series of images of basic limb muscle anatomy from dissections we’ve done of baby and adult Indian and White rhinoceroses. First, here’s what a rhino looks like underneath the skin:

But ahh that skin, that fabled “pachyderm” skin! A rhino’s greatest defense is also a real chore to get through in a dissection.  Here, we enlist the help of a crane and hook, hurrying to get down to the muscles of this forelimb before rotting takes over too much (as with other big animals, this is a tough race against time even in chilly England!):

Here is a closer look at that amazing armoured skin; sometimes 10cm or so thick:

Back to the forelimb muscles– stocky and well-defined for this athletic animal:

(late addition) Here are the massive shoulder muscles, such as the serratus and latissimus dorsi (this is a left limb in side view; head is toward the left):

And now a close look at the forearm muscles:

And then over to the hindlimb, here from an adult Indian rhino, whose thigh bone (femur) shows the characteristic giant “third trochanter” (toward the bottom centre of the image), which is an expanded bony attachment for the giant “gluteobiceps” muscle complex that retracts the femur for the power stroke in locomotion. Also, this specimen showed fascinating anatomy that I’d never seen before: the third trochanter has a thin bar of bone that extends up (toward the bottom left in the image) to fuse with the greater trochanter, opposite the head of the femur (upper left corner):

Damn my photography skills, cutting off the edge of that image and instead giving a view of my boots! Anyway, another interesting feature of that femur: the medial (inner) condyle of the femur (knee joint surface) has a pink stripe of worn cartilage. This is indicative of at least a moderate stage of arthritis, shown here (look for the pinkness amidst the shiny, healthy white cartilage on the upper right side). It is an exemplar of serious welfare problems that some captive, and probably some wild as well, rhinos face:

(late addition) Back up the limb, this baby White rhino shows the massive thigh muscles, especially that “gluteobiceps” that attaches to the third trochanter, noted above, and also showing the hamstrings:

Moving down the limb, we encounter the glorious three-toed perissodactyl foot of rhinos, and the robust hooves/nails, which are reasonably healthy in this animal– unlike others I’ve seen:

And the sole of that foot, showing a fairly healthy pad, below. Toward the rear (away from the nails), it culminates in a modest-sized fat pad, or digital cushion, akin to that in elephants but far less well developed and lacking the false “sixth toe” (predigit) (see also CT scan movie of the hindfoot above):

Here’s a view inside that marvelous foot, showing the HUGE digital flexor tendons. These help support the toes against gravity and, in theory, can act to curl them up– although in a rhino’s foot, as in an elephant’s, the toes are more like a single functional hoof, with reduced independence compared to a carnivore or primate:

And that ends our tour of rhinoceros limb anatomy and function. Help spread the word of how precious and threated rhinos are; educate yourself and others! And if you overhear someone talking about using rhino horn for medicine, try to politely educate them on the utter fallacy of this tradition. It is this cruel, greedy, ignorant practice that needs to die; not rhinos. I don’t enjoy receiving dead rhinos, on a personal level, even though the science excites me. I’d rather have many more alive and living good, healthy lives. And my team is trying to do what we can to help others on the “front lines” of rhino conservation make that happen.

For example, Will Fowlds, vet and co-owner of Amakhala Game Reserve, South Africa, recently sent us some images of a white rhino that had been caught in a poacher’s foot snare some years ago. The poor rhino still was having problems healing– we inspected x-ray images and external photos and helped to make an initial diagnosis of osteomyelitis, a nasty infectious, inflammatory foot bone/joint disease. We are following this case to hope that the rhino recovers and contribute help where we can, but the tough job belongs to the keepers/vets on the ground, not to mention the rhinos…

Furthermore, we’ve done foot pressure research covered here, and here is an example of the data we’ve collected (image credit: Dr Olga Panagiotopoulou), showing high pressures on the toes and low pressures on the foot pads:

Big thanks to people on my team that have helped with this and related research: Dr Olga Panagiotopoulou (and Dr Todd Pataky at Shinshu University, Japan), Dr Renate Weller in the VCS Dept at the RVC, Liz Ferrer at Berkeley, and former undergraduate student researchers Sophie Regnault, Richard Harvey, Hinnah Rehman, Richard Sheehan, Kate Jones, Bryony Armson and Suzannah Williams.

A White rhino’s heart, with more images below, all courtesy of William Perez’s Veterinary Anatomy Facebook pages. A mass of around 10kg (22 lbs weight) is not unusual! (Compare with even larger elephant heart)

White rhino closeup: coronary arteries

White rhino: branches of left coronary artery

White rhino heart: right atrium

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Here now is the promised blog post, which uses the rhino foot mystery pic as a springboard to address a phenomenon that is a bit better known, partly because it is an even worse situation and involving (arguably) even more charismatic critters: elephants.

A rotating movie of a CT scan reconstruction is a good way to kick this off:

This shows the right hind foot of an Asian elephant that had mild pathology; mostly a roughening of some of the bone surfaces that is called osteitis (proliferative bone growth possibly due to infection or other irritation) and perhaps a mild case of degenerative joint disease such as osteoarthritis. But this is nothing compared to the severe cases we’ve observed in other elephant feet, and indeed may not have anything to do with why this elephant died (I’m not sure; I was given very little medical history for this one).

If you want more elephant anatomy lessons, see the videos from the posting on six-toed elephants. I will proceed assuming some basic familiarity with bones of the feet in animals, although you may be just fine even without that.

About 50% of elephants in captivity die from foot disorders of one kind or another. Elephant keepers spend a huge amount of time and energy taking the best care of elephant feet that they can, but a variety of factors including anatomy, biomechanics, exercise, obesity, ground surface, hygiene, “hoof” care including trimming, nutrition, and much more are part of the very complex causal nexus underlying these disorders. Wild elephants get similar problems, too, but less frequently (e.g. in drought periods, I’m told); there are few solid data on this, however.

Onwards, then! I shall present a cavalcade of horrific examples of the kinds of elephant foot pathology that we have observed in specimens that have come through my freezers at the RVC.

Let’s start with what one of our vets might see on examination of a live elephant at a zoo:

This is an x-ray image of the third (on the left) and fourth (on the right) toes of an elephant’s front foot. The RVC (Dr. Renate Weller and myself) have developed protocols to take such x-rays on live elephants.  The anatomy shown here is pretty normal and non-pathological. So with that in mind, check this out; toes four (on the left) and five (on the right), different animal:

Ouch! Digit 4 (“ring finger”) has a proliferation of bone that is characteristic of an animal with osteomyelitis: a flowering of bone in response to infection and painful swelling, probably caused by an abscess on the toe’s sole/nail. This animal was put down because of its unresolvable misery from this disorder. Oddly, we see toe 4 as well as 3 and 5 as the most commonly pathological; toes 1 and 2 seldom are. We’ll be discussing this in a new paper coming out soon; I’ll get back to that another day.

Assuming such conditions don’t resolve, the next place the foot may end up is in The Freezer at the RVC, and then into our CT scanner before we do postmortem dissections and a report on the pathologies so the zoo knows what went wrong. Here’s an example of what we cut off the end of the fourth toe of such an animal:

Just looks like a glob of tissue, right? The joint between two segments of the toe is visible as a pinkish white structure on the right side, with some bleeding on the cartilage where it wore down to the bone surface. But it gets worse. Here is how that same toe bone looked when we cleaned it up (boiling and bleaching away soft tissues):

Here, that same roughened joint surface is visible at the top of the specimen. Two toe bones have become fused together (the bottom one is not visible), encased in a cocoon of lacy, spiny bone. Again, ouch. The next specimen had a different kind of “ouch”- its fifth toe basically shattered:

That toe is almost unrecognizable, having disintegrated rather than proliferated its bony scaffolding. Other specimens may be in less extreme states of pathology but still likely to have been in pain:

The label here says it all; third toe with a cyst where an infection entered the bone.

This one, the end of a third metatarsal, shows degenerative joint disease with a loss of articular cartilage, and holes where abrasion has worn down into the bone and caused bleeding. In contrast, and to give you a breather from the horrors, here is a healthy, younger elephant’s similar joint surface:

Nice white, fresh, shiny cartilage! Ahhh…

But then we dive back into Grand Guignol-level aberrations:

Here we’re looking at the back side of a right hind foot of an elephant, at the level of the ankle joint. The joint capsule surrounding the ankle joint has been cut open in my dissection to expose the terribly pathological, but still somewhat white and shiny, cartilages (middle of the image) which have been abraded (in some regions) but also extended by new bone formation (in other regions) to creep around the back of the ankle. Here, the bone growth was fulfulling a role to limit joint mobility and thereby restrict painful joint motions- the joint was fusing into an ankylosis (no, not an ankylosaur,  but same Greek root). Here is a closer look, removing the tibia and fibula that were at the top of the screen in the above image, and looking down onto the ankle joint surface:

You should be able to more clearly see how the cartilage and underlying bone are not forming a smooth edge, as they should on the talus (ankle bone), but rather an irregular, jagged contour (area to the right of the label). This animal would have been visibly lame, to say the least; elephant ankles can’t move much even in normal animals but this one was even less mobile. We’ve had some specimens where the ankle was so fused it was totally immobile and took a saw to separate the two sides of the joint. Oddly, I haven’t seen an ankylosis like that in the wrist, which in normal elephants is as flexible a joint as the ankle is inflexible.

Pathologies like these sadly aren’t uncommon in elephant feet but zoo/park keepers are doing their best to turn the trend around. Zoo conditions generally were a lot worse 50 years ago. The pictures below document this, from museum specimens we’ve studied (among many others) at the University Museum of Zoology at Cambridge and the Natural History Museum in London. See what pathologies you can spot! Some are from wild-shot animals, reinforcing that foot pathologies are not just a zoo thing. (click to embiggen)

      

Zoo/park conditions are improving now– in the UK for example, elephants are being moved into more safari park-like environs and given more varied surfaces to walk on or even dig (e.g. sand at Chester Zoo). But because elephants live long lives, and foot pathologies sometimes cannot be reversible (or even detectable, sometimes), any pathologies existing now may well still be evident, or even worsen despite the best care, for decades to come. The lag time for fixing the global problem of elephant foot pathologies is not a short one. I won’t get into the controversy over whether elephants should be in zoos/parks or not, but at least for the short to medium term they are, and we need to make the best of that. The images in this post help show why, and perhaps point a way toward how.

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I will go into more detail soon on the broader subject that this involves, but am posting this image as a teaser– what’s up with this foot from my freezer?

Other than the obvious dead-ness and non-attached-to-body-ness…

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Well, it’s time for the Grand Reveal of what the picture in the previous post is (reproduced below). The guesses ranged from bird to wallaby/kangaroo to the stuff of nightmares. And indeed Nick Gardner got it right first, it is a wallaby. Specifically, it is a Bennett’s Wallaby (Macropus rufogriseus rufogriseus), which is the Tasmanian island form related to the Red-necked Wallaby, and an animal that has gone feral (along with mara and other cool critters) in Whipsnade Zoo near the RVC. You can tell it is a wallaby and not a bird, because there is an “Achilles tendon” attaching to a calcaneal tuber (“heel bone”) on the back side of the limb (shown with asterisk below) that birds lack, and if you look closely the toes are hairy, lack bird-like claws, and a few other details like the profile of the musculature are very different; more mammalian than avian. The stump of the muscular tail (cut off) is also a clue. Although the avian similarity in the case of wallabies is still striking, which is one reason I chose this image. Well done, Nick!

I found this picture in my archives and remembered when it was taken back in ~2005- some lab members received some frozen wallaby legs and thawed them out to use in experiments. They tried to compress the legs in an Instron machine (mechanical testing system; partly visible at the top of the pic) to see what the passive, springlike properties of the legs are like in a wallaby, vs. the properties they could measure in a living animal. (The shiny white reflective areas in the pic are for tracking joint motions) And I thought it was a freaky cool pic, so I shared it.

I also posted that pic because my team has done some in vivo analysis of the leg properties in such animals (previous news story here; paper in preparation), and because we use this technique of loading cadaveric legs in such machines quite routinely. We did this for elephant feet to study how the “sixth toe” of elephants works, and we’re analyzing data (as I write) for how elephant feet and rhino feet deform or move when loaded similarly. This method has a long history; we didn’t invent it; perhaps most famously used for studying horse limb mechanics [pdf example], which have a lot of passive properties (almost everything below the elbow/knee is non-muscular). Many animals’ limbs are tendinous/elastic toward their distal end (toes), so the limbs tend to become less actively controlled by the nervous system and become more of a mechanical control system (sometimes involving a non-neural “preflex“) in that region; although it’s all a matter of relative degree of passive:active control in different situations, species, and limbs.

The picture below shows an x-ray of an elephant’s whole hind foot, in which you should be able to see the bones (brighter white) of the foot surrounded by a lot of soft tissue, mostly more passive kinds like fat, skin, fascial sheets, ligaments and tendon.

Here (further below) is a preliminary image from our elephant foot studies in progress, intended to reveal the passive properties/motions of the feet so we can figure out how those properties are combined with more active control, and how actively elephants control their feet vs. other, possibly more ‘passive-footed’ animals like horses. This is interesting from anatomical and evolutionary perspectives, and for helping with foot health problems that are serious concerns for such animals– more about that later. The arrow in the picture below shows where a lot of the motion is: at the knuckle (metatarsophalangeal) joints of the toes; the rest of the foot tends to rotate around these mobile joints. We can’t peer inside living elephant feet to see if they actually do this, but we can compare the external motions, pressure patterns, and other data from living and dead elephant feet to see how they match up, which is what we’re doing now — and we’re doing the same thing with rhinos, which have cool 3-toed, more “hoof-like” feet, as opposed to the 5-toed, fatter feet of elephants. To get this image, we’ve had to put the foot inside a custom-made device using a car jack to apply a constant load, and a wooden framework to hold the specimen still, and then run it through a CT machine in unloaded and then loaded states to see how the bones move. Here is what the crazy apparatus looks like, with enthusiastic undergrad for scale:

This, below, is a right hind foot (pes) of an Asian elephant, shown from the inside of the foot (toes are numbered 1-4 from the big toe/hallux toward the outside of the foot; 5th toe is not visible). The yellow image is the relaxed, unloaded foot; the green is after applying a large load equivalent to the animal standing on one foot (or running quickly). Notice how the third metatarsal (the long bone that the arrowhead is touching) for the unloaded state is in front of that for the loaded state, whereas yellow and green images of the bones toward the tip of the toes are overlapping more, indicating they did not move much/at all. That tells us that the motion is occuring at the joint indicated (“knuckle”), which makes sense anatomically, because that joint looks like it has a lot of mobility.

Image

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In case you missed the story about this paper released just before Xmas, here are some links to stories about “From flat foot to fat foot: Structure, ontogeny, function and evolution of elephant “sixth toes,” in Science, 2011:

1. The paper (free download from my publications list; nice policy, Science!)

2. Our website about the paper (more imagery goodness!)

3. Ed Yong’s first (Nature News) and second (more detailed blog) article

4. BBC News’s story

5. Reuters TV‘s excellent video

6. Science Now/Wired’s story

7. Daily Mail‘s story (not a daily fail, in this case)

Clarification: it’s not a real sixth toe in elephants; it’s a false, toe-like structure (“predigit”) made from other tissue. That confusion seeped into some media stories. But this whole story ties into the thorny question of what a digit (finger/toe) is and how we can tell (e.g., notions of homology). Regardless, the elephant predigits are present in all four feet, and are super duper cool!

Most importantly for this blog, that research relied, and still relies, on our fabulous freezers to keep the elephant “toes” in snuggly cold conditions until we wanted to study them.

The research is continuing- I’ll post more about that later. We’ve been doing lots more histology to explore the complex ways that these predigits are formed, and also studying how they function (ex vivo) in more 3D detail than before (with new comparisons to rhino feet). Also, a new paper of ours will come out in J Experimental Biology very soon. It elaborates on how whole elephant feet function, across ontogeny, using in vivo pressure patterns.

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