2015 AAPA conference: More brain growth

The American Association of Physical Anthropologists is holding its annual meeting next year in St. Louis, in my home state of Missouri (I’m from Kansas City, which is by far the best city in the state, if not the entirety of the Midwest). I’ll be giving a talk comparing brain size growth in captive and wild chimpanzees, on Saturday 28 March in the Primate Life History session. Here’s a sneak peak:

Velocity curve for brain size from birth to 5 years in wild (green) and caprive (blue) chimpanzees. For the captive models, the dashed line is fit to the raw brain masses, and the solid line is fit to the estimated endocranial volumes.

Velocity curves for brain size growth from birth to 5 years in wild (green) and caprive (blue) chimpanzees. The wild data are endocranial volumes, but the captive specimens are represented by brain masses. So the captive data are modeled for both the original masses (dashed) and estimated volumes (solid). Wild data are from Neubauer et al. 2011, captive data from Herndon et al., 1999.

Abstract: This study compares postnatal brain size change in two important chimpanzee samples: brain masses of captive apes at the Yerkes National Primate Research Center, and endocranial volumes (ECVs) of wild-collected individuals from the Taï Forest. Importantly, age at death is known for every individual, so these cross-sectional samples allow inferences of patterns and rates of brain growth in these populations. Previous studies have revealed differences in growth and health between wild and captive animals, but such habitat effects have yet to be investigated for brain growth. It has also been hypothesized that brain mass and endocranial volume follow different growth curves. To address these issues, I compare the Yerkes brain mass data (n=70) with the Taï ECVs (n=30), modeling both size and velocity change over time with polynomial regression. Yerkes masses overlap with Taï volumes at all ages, though values for the former tend to be slightly elevated over the latter. Velocity curves indicate that growth decelerates more rapidly for mass than ECV. Both velocity curves come to encompass zero between three and four years of age, with Yerkes mass slightly preceding Taï ECV. Thus, Yerkes brain masses and Taï ECVs show a very similar pattern of size change, but there are minor differences indicating at least a small effect of differences in habitat, unit of measurement, or a combination of both. The overall similarity between datasets, however, points to the canalization of brain growth in Pan troglodytes.

Kazakhstan’s killer cats

I’m reading up on previous paleontological research that’s taken place here in Kazakhstan, planning for future work. There aren’t any human fossils known from here (at least, none to my knowledge). But, I did stumble upon this badass, sabre-toothed cat from the Late Miocene (over 5 million years ago):

From Sotnikova, 1992. Original caption: Fig. 2. Machairodus kurteni, Kalmakpai (PIN-2433/287), skull, ventral and lateral view.

From Sotnikova, 1992 (mandible not shown). Original caption: Fig. 2. Machairodus kurteni, Kalmakpai (PIN-2433/287), skull, ventral and lateral view.

The skull was described by MV Sotnikova in 1992, and comes from a site called Kalmakpai in the Zaysan Basin in East Kazakhstan. For perspective, Sotnikova says the skull is about the same size as an adult African lion. This is much larger than the wildlife I’ve seen lately in snow-soaked Astana (I trailed a large white rabbit in Presidential Park by the river on my run today. Not as badass).

This is a reminder that the Big Cats once had a much larger geographical distribution than they do today. The skull above belongs to the genus Machairodus, which is also known from Africa, Europe and North America. Machairodus is closely related to Homotherium, another large, geographically dispersed genus of sabre-toothed cat from the Pliocene (including at Dmanisi).

Of course, extinction isn’t exclusive to the deep past: the Caspian Tiger used to roam parts of southern Kazakhstan and other areas of Central Asia, going extinct only in the past few decades.

Results of the toe-tally easy lab activity

Alternate title: Dorsal canting in primate PPP4s

Earlier this year I suggested a classroom activity in which students can scrutinize the evidence used to argue that the >5 million year old (mya) Ardipithecus kadabba was bipedal. To recap: Ar. kadabba is represented by some teeth, a broken lower jaw, and some fragmentary postcrania. The main piece of evidence that it is a human ancestor and not just any old ape is from a single toe bone, and the orientation of its proximal joint. In Ar. kadabba and animals that hyperdorxiflex their toes (i.e., humans and other bipeds when walking), this joint faces upward, whereas it points backward or even downward in apes. This “dorsal canting” of the proximal toe joint has also been used as evidence that the 4.4 mya Ardipithecus ramidus and 3.5 mya owner of the mystery foot from Burtele are bipedal hominins. A question remains, though – does this anatomy really distinguish locomotor groups such as bipeds from quadrupeds?

Use ImageJ to measure the canting angle between the proximal joint and plantar surface. Proximal to the right, distal to the left.

STUDENT SCIENTISTS TO THE RESCUE! Use ImageJ to measure the canting angle between the proximal joint and plantar surface, as I’ve done on this Japanese macaque monkey (they are not bipedal). Proximal to the right, distal to the left Note I changed the measured angle from the March post.

I sicked my students in Ant 364 (Human Evolutionary Developmental Biology) here at NU on this task. I had students look at only 11 modern primates from the awesome KUPRI database. Most groups are only represented by 1 (Homo sapiens, Hylobates lar and Macaca fuscata) or two (Pongo species and Gorilla gorilla) specimens, all adults. For chimpanzees (Pan troglodytes) there is one infant and four adults. The database has more individuals, and it would be better to include more specimens to get better ideas of species’ ranges of variation, but this is a good training sample for a class assignment. The fossil group includes one Ardipithecus ramidus, one Ar. kadabba, one Australopithecus afarensis, and the PPP4 of the mystery foot from Burtele. The human and all fossils except Ar. kadabba are based off of lateral photographs and not CT scans like for the living primates, meaning there may be some error in their measurements, but we’ll assume for the assignment this is not a problem. Here are their results:

Dorsal canting angle of the fourth proximal pedal phalanx in primates.

Dorsal canting angle of the fourth proximal pedal phalanx in primates. The lower the angle, the more dorsally canted the proximal joint surface. The “Fossil” group includes specimens attributed to ArdipithecusAustralopithecus and something unknown.

Great apes have fairly high angles, meaning generally not dorsally canted proximal joint surfaces. The two gorillas fall right in the adult chimpanzee (adult) range of variation, while chimp infant and orangutans have much higher angles (≥90º means they’re actually angled downward or plantarly). The gibbon (Hylobates) is slightly lower than the chimpanzee range. The macaque has an even more dorsally canted joint, and the human even more so. The fossils, except the measurement for Ar. ramidus (see note above), have lower angles than living apes, but higher than the human and the monkey. If dorsal canting really is really a bony adaptation to forces experienced during life, then the fossil angles suggest these animals’ toes were dorsiflexed more so than living great apes (but not as much as the single monkey and human).

This lab helps students become familiar with CT data, the fossil record, taking measurements (students also measure maximum length of the toe bones and look at the relationship between length and canting), analyzing data, and hypothesis testing. You can also have fun exploring inter-observer error by comparing students’ measurements.

Here’s the full lab handout if you want to use or modify it for your own class: Lab 5-Toe instructions and report

Lessons from limb lab (activity)

This semester I have added a lab component to my Introduction to Biological Anthropology class. Lab activities and assignments provide students with opportunities to gather data, to think about them in the context of various theories, and to learn about how to analyze them. This past week’s lab looked at limb proportions within our classroom, in the context of “Allen’s rule” – in colder climates, animals tend to have relatively shorter distal limbs (radius+ulna and tibia+fibula). Allen’s rule, along with “Bergmann’s rule,” describe ecogeographic variation in humans and other animals: body size (i.e., mass) and shape (i.e., limb proportions) tend to vary with climate, such that populations living in colder environments tend to have less surface area relative to body mass, as an adapation to retain body heat.

A simple and effective way to quantify the relative lengths of limbs is through ratios: in this case we examined the crural and brachial indices. Here I’ve plotted average human indices against latitude as reported in a recent paper by Helen Kurki and colleagues (2008):

Left: Crural index (tibia length/femur length) related to latitude, as a proxy for climate. Right: Brachial index (radius length/humerus length) plotted against latitude. Data from Kurki et al. (2008). Black=female, red=male. Dashed lines are  regression slopes for each sex, and solid lines indicate the 95% confidence limits of the regression lines.

Human limb proportions related to latitude, as a proxy for climate. Left: Crural index (tibia length/femur length). Right: Brachial index (radius length/humerus length). Higher indices mean relatively longer tibia or radius. Data from Kurki et al. (2008). Black=female, red=male. Dashed lines are least squares regression lines for each sex, and solid lines indicate the 95% confidence limits of the regression lines. How well will our class’s data fit these models?…

These plots are consistent with Allen’s rule – the distal limb segments become relatively shorter with increasing latitude. In lab, we test whether our limb proportions reflect this presumably ecogeographic pattern. Here in Astana we are at 51ºN latitude, so these regressions predict that our class should have crural indices between 0.82-0.84, and brachial indices between 0.72-0.79. How well does our class fit these model’s predictions?

Pretty terribly.

…Pretty terribly. Plots are same as above, except with our class’s data added at 51º N latitude (vertical lines). In each, the vertical lines span the class’s 95% range (black=females, red=males), with the dots marking each sex’s average. Kazakhstan is huge, and students could have grown up in latitudes from 42º-55º N, but even assuming students had all come from Shymkent their distal limbs still appear much longer than expected.

These plots show that students in the class have longer distal limbs than expected – both for our latitude, and for humans generally. The poor fit of my students’ limb proportions probably doesn’t mean they’re bad humans. Instead, we probably deviate because we compared apples to oranges: the Kurki data were dry long bones measured on an osteometric board, whereas I had my students do their best to palpate and measure the maximum lengths of their own bones beneath layers of fat, muscle, skin, and clothing. Our high indices probably reflect the underestimation of humerus and femur lengths, whose most proximal points that can be palapated (greater tubercle and trochanter, respectively) lie a bit lower than the respective heads, which would have been included in the Kurki measurements.

It was interesting to review these plots with students. Even though they’re fairly new at reading graphs like these, there was an audible gasp and bewildered muttering when their own data went up on the board. I myself was surprised at these results, but I’m happy with how the exercise went. This particular ‘study’ helps students learn about ecogeography, adaptation and human variation, as well as the importance of homology and comparing like with like.

eFfing Fossil Friday: Funky facial flanges #FFF

David Reich and colleagues announced in this week’s Nature the discovery of a new species of extinct mammal, Vintana sertichi, that lived in what is now Madagascar between 66-72 million years ago. The species is based on a very well-preserved cranium of an early gondwanatherian (if you want to impress your friends this weekend, gratuitously use the word “gondwanatherian”). I don’t know much about early mammals like this, but it sounds like it was a weird creature (see the Stony Brook press release). Just looking at it’s face there’s something that sticks out as strange:

Ventana sertichi cranium (Reich et al. 2014, Figure 1a). Left is a 3D CT reconstruction, right is a line drawing highlighting all the individual bones (so many cranial bones). The view is from the right side, so the nose is on the right, the eye is the big hollow in the middle, and the back of the skull is on the left. The jugal flanges are the downward projections.

Vintana sertichi cranium (Reich et al. 2014, Figure 1a). On the left is a 3D CT reconstruction, and on the right is a line drawing highlighting all the individual bones (so many cranial bones). The view is from the right side, so the nose is to the right, the eye socket is the shadowy hollow in the middle, and the back of the skull is on the left. The jugal flanges are the downward projections.

Jutting downward from the sides of the jaw are ‘jugal flanges,’ projections of bone on the homologs of human cheeks. Projections of like these usually serve as muscle attachment sites, and the size of the projection generally reflects the size of the muscle. These facial flanges anchor the masseter muscle, a major chewing muscle that helps close the jaw. The size of this flange in Vintana suggests its chomp packed a punch. A debilitating bite. A face not even a mother could love (so now they’re extinct).

Vintana‘s bony tear-catchers caught my eye because most primates I’ve seen have, you know, less heinous faces. Scouring the internet, big jugal flanges are a fairly rare sight, but can apparently be found in glyptodonts (giant, armadillo-like mammals that lived tens of thousands of years ago) and various sloths. The closest thing I’ve seen to this gross bony flange in Primates are on the zygomatic bones of some extinct, baboon-like animals, such as Dinopithecus ingens:

Fragmentary skull, viewed from the top, of Papio (a.k.a. Dinopithecus) ingens, from Swartkrans, South Africa. Photo credit: CalPhotos.

Fragmentary skull, viewed from the top, of Papio (a.k.a. Dinopithecus) ingens, from Swartkrans, South Africa. As a punishment for its zygomatic excess, its face was confiscated. Photo credit: CalPhotos.

and Theropithecus brumpti

Theropithecus brumpti from the Omo basin. Photo credit: CalPhotos.

Theropithecus brumpti from the Omo Basin, Ethiopia. Photo credit: CalPhotos.

So some primates dabbled in jugal flangery like Vintana, but Natural Selection was having none of it. Anyway, Vintana overcame this craniofacial adversity with characteristic Mesozoic moxie, and is an important piece in the puzzle of mammal evolution. It will be interesting to see what other mammalian surprises the Mesozoic has in store for paleontologists.

Can ‘ape-like’ actually be ‘human-like’?

I’m reading up on life history in Homo erectus for a few projects I’m working on, and something’s just caught my eye. A 2012 issue of Current Anthropology presents a series of papers from the 2011 symposium, “Human Biology and the Origins of Homo.” This issue is full of great stuff, and to top it all off, it can be accessed online for free! (here’s the JSTOR link)

Gary Schwartz has a paper here recounting what is known (or as he stresses, what is still largely unknown) about growth and life history in early Homo. Dental evidence accumulated over the past 30 years has pointed to a rapid (ape-like) life cycle for fossil hominins, in comparison with a slow, long and drawn out human pattern. But much of the evidence against a human-like pattern is somewhat indirect. For instance, Holly Smith (1991) has shown that there’s a pretty tight relationship between brain size and age at first molar (M1) eruption in Primates:

M1 crancap

Fig. 1 from Schwartz (2012). “Bivariate plot of ln M1 emergence age in months (y) versus ln cranial capacity in cubic centimeters (x) for a sample of anthropoids.” The hominins and humans are the open shapes, to which I’ve visually fitted the red line.

It’s a very high correlation (r=0.98). This means that armed with simply an animal’s cranial capacity, which is fairly easy to estimate given complete enough fossils, one can estimate with a bit of confidence its likely age range for M1 emergence. With brain sizes between apes’ and ours, fossil hominins can be estimated to have erupted their M1s at younger ages than us. Many subsequent studies of tooth formation, based on the microscopic remnants of tooth development, have supported these inferences. So presumably, faster, ape-like dental development could be extrapolated to mean ape-like body growth rates and other aspects of life history as well.

But although this is a tight relationship, there are deviations. As Schwartz notes in the article, and others have noted before, high correlations found when examining large interspecific groups (e.g., primates as a whole) often break down when the focus is on smaller groups of more closely related species (e.g., just apes). Based on the relationship figured above, humans are expected to erupt M1 around 7 years of age, but nearly all humans erupt M1 closer to 6 years (hence the open diamond for humans is below the regression line). What hominins appear to share in common with humans is a younger age at M1 eruption than expected for primates of their brain sizes (the red line I’ve added to the figure).

Hominins’ faster dental development and eruption may be ape-like in absolute terms, but eruption ages may be human-like when their brain size is taken to account. As with many life history variables, the significance of this similarity (if anything) is difficult to ascertain.

eFfing Fossil Friday: Frozen Femur

A 45,000 year old human femur from Siberia provides new information about genetic mutation rates and modern human origins. As Quiaomei Fu and colleagues report in this week’s issue of Nature, this seemingly simple leg bone carries so much information, not because of its gross anatomy, but because of the ancient DNA it preserves.

The femur wasn’t discovered by paleontologists, but by an artist/historian looking for fossils around the Irtysh River. The bone came from from a site called Ust’-Ishim, only some 650 km north of the snowy capital where I work in Kazakhstan:

Ust'-Ishim

The site in question, Ust’-Ishim is marked by the yellow star. The red and blue sites to the southeast are other Upper Paleolithic sites. Okladnikov (3) and Denisova (4) have also yielded fossils preserving ancient DNA. Modified from Fu et al. figure 1.

The bone was directly radiocarbon dated to around 45,000 years ago. With a fairly precise age of the bone, Fu et al. could estimate the rate at which genetic mutations arise, by counting the number of new mutations in recent humans that aren’t shared by the Ust’-Ishim femur. This led to an estimate of around 0.43×10−9  new mutations per site per year. This is a relatively low rate compared to estimates based on geologically older fossils, but consistent with more recent estimates that directly compare parents and offspring.

The Ust’-Ishim individual had levels of Neandertal ancestry comparable to living Eurasians (~2.3% of the genome), but there is no evidence of any Denisovan ancestry. Because this individual lived closer to the date of modern-Neandertal admixture, the Neandertal segments of its genome are longer than in modern people (recombination over generations breaks these regions apart into shorter segments). Knowing about recombination rates, Fu et al. could infer that admixture between Neandertal and modern human populations occurred between 50-60,000 years ago.

This eFfing Friday fossil provides more tantalizing evidence for DNA-bearing human fossils just across the Kazakhstan border. With Ust’-Ishim to the north, Denisova and Okladnikov caves to the east, and Teshik Tash to the south, my colleagues and I are very keen to find similar sites here on the KZ side.

Reference: Fu et al. 2014. Genome sequence of a 45,000-year-old modern human from Siberia. Nature 514: 445–449. doi:10.1038/nature13810.