john hawks weblog

paleoanthropology, genetics and evolution

Sahara at least 7 million years old

Sat, 2006-02-11 12:07 -- John Hawks

A concise 4-paragraph article by Mathieu Schuster and colleagues reports on dune deposits that show the Sahara formed during the Late Miocene.

After the mid-Holocene humid period (6000 years ago), arid conditions developed throughout North Africa, culminating in the formation of the Sahara, which is the largest warm-climate desert on Earth (9,000,000 km2). However, earlier desert recurrences in the region are also documented. Direct evidence for eolian deposition is given by thermoluminescence dating for the Late Pleistocene; e.g., in Mauritania [25 to 15 thousand years ago (ka)] (1) or in Tunisia (86 ka) (2). The latter is currently considered as the oldest terrestrial record for desert conditions in the Sahara (2), even if firm evidence exists for a pre-Quaternary Great Western Sand Sea in Algeria (3). Some earlier arid episodes (Miocene-Pliocene) were also suggested by marine records off West Africa (4); but until now, no contemporary in situ eolian deposits were known in the Sahara region. In the northern Chad Basin, we recently identified and dated widespread outcrops of eolian dune deposits that are distributed over an area more than 2000 km2. Our results testify that the onset of recurrent desert conditions in the Sahara started at least 7 million years ago (5-7) (Schuster et al. 2006:821).

The desert comes and goes, expanding and contracting -- and those vacillations are recorded by this earliest evidence, also:

In the Toros Menalla region, these eolian sandstones are conformably overlain by a horizon bearing abundant vertebrates fossils, including Sahelanthropus tchadensis, the earliest known Hominid [sic] (5, 7). In this horizon, named the Anthracotheriid Unit, biostratigraphic correlation of the mammalian fauna indicates an age of 7 Ma (5–7).

Now, this isn't news (which I'm sure Science didn't bother to check) since Vignaud and colleagues (2002) published the same evidence, complete with the wind direction chart:

The lower part of the section (at least 4 m thick) is composed of fine to very fine white sands, poorly cemented, and is mainly constituted by numerous quartz grains, without matrix. The grains are well sorted, well rounded, matt and frosted, and are strong evidence for aeolian modelling. The foreset laminations (avalanche laminations in front of the aeolian dune) represent a typically aeolian deposit. These sands show cross-beddings that progressively decrease in size from the bottom (1 - 2 m) to the top (20 cm). This facies exhibits typical alternations of grain-fall and grain-flow laminations, characteristic of aeolian dune deposits. Our interpretation is confirmed by frequent wind ripples at the foot of the fossil dunes, whose crests are perpendicular to the direction of dune progradation. These fossil dunes are, to our knowledge, the oldest evidence for desert conditions in the southern Sahara area (Vignaud et al. 2002:152).

I guess this is the science journal equivalent of getting "punk'd" -- "Ha ha! You published what we printed four years ago!"

I opened up the Vignaud paper to double-check the paleoenvironment in the fossil-bearing layer. From the faunal list, they conclude this:

The oldest known East African hominids (Ororrin [sic], Ardipithecus) are contemporary with faunas associated with wooded environments. Younger australopithecines lived in a wider range of habitats. In contrast, the TM 266 vertebrate fauna contemporary of the Toros-Menalla hominid suggests a mosaic of environments from gallery forest at the edge of a lake area to a dominance of large savannah and grassland. Determining the precise habitat of the TM 266 hominid locality among the mosaic of environments available to it constitutes a research challenge to be met by further laboratory and field studies currently in progress (Vignaud et al. 2002:155).

They (Vignaud et al. 2002) interpreted the succession of dune and lacustrine deposits to mean that the hominids lived in a mosaic environment near sandy desert, but locally including marshy/swampy, lake, and gallery forest. An alternative interpretation might be that the desert really receded (or disappeared) during the later time period when the hominids were there. In either case, the paleoenvironment is interesting, because it means that the Sahelanthropus-like primates colonized (and possibly repeatedly recolonized) areas that were periodically dune desert (and therefore probably not habitable by large primates). This may not mean much in terms of locomotion -- the hominid-bearing unit is clearly water-rich, and we can't refute the idea that the surroundings were as woodland-like as those preserved in the Late Miocene Middle Awash localities.

But I think it is a good hypothesis that all of these apes (or hominids) were very cosmopolitan compared to extant chimpanzees and gorillas. The question is whether their actual dispersal abilities were different from chimpanzees. Prehistorically, genetics would seem to indicate that chimpanzees had long-distance dispersal; the only fossil evidence of chimpanzees has been found in a region that historically did not support chimpanzees; and they today successfully utilize relatively open savanna at the eastern end of their range.

So it is by no means obvious that the cosmopolitan nature of these Late Miocene lineages would have required a specialized terrestrial adaptation -- at least not beyond the specialization of knuckle-walking. So why become bipeds?

References:

Schuster M et al. 2006. The age of the Sahara Desert. Science 311:821. Full text (subscription)

Vignaud P et al. 2002. Geology and paleontology of the Upper Miocene Toros-Menalla hominid locality, Chad. Full text (subscription)

PaleoAnthropology going open-access

Sat, 2006-02-11 11:54 -- John Hawks

I got a newsletter from the Paleoanthropology Society today, and it included this:

In late 2005 the Society's contract with the University of Pennsylvania Press expired. The Officers and Editors have decided to take over the responsibility of publishing our journal, PaleoAnthropology, with shared support from the University of Pennsylvania Museum. Because of this, a number of changes will take place.

The most important of these is that, beginning immediately, the journal will be accessible free of charge to everyone, including non-members of the Paleoanthropology Society. The journal can be accessed at its own address (http://www.paleoanthro.org/journal/default.htm) or through the main page of the Society's website (http://www.paleoanthro.org/).

With the advice of the Board of Editors, we will also be adding additional features to the journal. Currently the journal has been devoted to the publication of articles, book reviews, and the abstracts of the annual meetings of the Society. This will be expanded, in part, to include commentaries on articles, summaries of current work in the various fields of paleoanthropology, and other features.

I think this is super cool, and I can't overstate the importance of this in my decision about where to send research. Nothing galls me more than having to stick that "subscription only" reminder next to things I link to, because I know there are a lot of interested people who can't get them.

And this journal is especially great because it allows the distribution of datasets, color illustrations, and now anybody can get it!

A Darwin Day parable

Sat, 2006-02-11 09:09 -- John Hawks
"We're going to arm you with Christian Patriot missiles," Ham, 54, recently told the 1,200 adults gathered at Calvary Temple here in northern New Jersey. It was a Friday night, the kickoff of a heavily advertised weekend conference sponsored by Ham's ministry, Answers in Genesis.

There has been a lot of attention this week to the congregations that are observing Darwin Day, as a recognition that evolution and religion are not in conflict.

The LA Times has an article profiling Ken Ham, who would be on the, um, other side of this issue.

A former high-school biology teacher, Ham travels the nation training children as young as 5 to challenge science orthodoxy. He doesn't engage in the political and legal fights that have erupted over the teaching of evolution. His strategy is more subtle: He aims to give people who trust the biblical account of creation the confidence to defend their views -- aggressively.

He urges students to offer creationist critiques of their textbooks, parents to take on science museum docents, professionals to raise the subject with colleagues. If Ham has done his job well, his acolytes will ask enough pointed questions -- and set forth enough persuasive arguments -- to shake the doctrine of Darwin.

The article reports on one of these training sessions. Here's the part excerpted by Ann Althouse:

Evangelist Ken Ham smiled at the 2,300 elementary students packed into pews, their faces rapt. With dinosaur puppets and silly cartoons, he was training them to reject much of geology, paleontology and evolutionary biology as a sinister tangle of lies.

"Boys and girls," Ham said. If a teacher so much as mentions evolution, or the Big Bang, or an era when dinosaurs ruled the Earth, "you put your hand up and you say, 'Excuse me, were you there?' Can you remember that?"

The children roared their assent.

"Sometimes people will answer, 'No, but you weren't there either,' " Ham told them. "Then you say, 'No, I wasn't, but I know someone who was, and I have his book about the history of the world.' " He waved his Bible in the air.

"Who's the only one who's always been there?" Ham asked.

"God!" the boys and girls shouted.

"Who's the only one who knows everything?"

"God!"

"So who should you always trust, God or the scientists?"

The children answered with a thundering: "God!"

I notice the annual budget of Answers in Genesis ($15 million) is almost enough to sequence the genome of another mammal species -- every one of which shows 3 billion or so marks of evolution.

We need to get more fossils into schools.

Janet Monge, Darwinista

Thu, 2006-02-09 23:00 -- John Hawks

The AP is running an article about Darwin Day this Sunday, and Gretchen spotted it on MSNBC accompanied by a photo of Janet Monge!

Very cool! Here's some of the article:

PHILADELPHIA - Thanks to the "intelligent design" movement, Charles Darwin's birthday is evolving into everything from a badminton party to church sermons this weekend.

Defenders of Darwin's theory of natural selection are planning hundreds of events around the world Sunday, the 197th anniversary of his birth, saying recent challenges to the teaching of evolution have re-emphasized the need to promote his work.

"The people who believe in evolution ... really just sort of need to stand up and be counted," said Richard Leventhal, director of the University of Pennsylvania's Museum of Archaeology and Anthropology. "Evolution is the model that drives science. It's time to recognize that."

Human Genome Project afterglow

Wed, 2006-02-08 23:35 -- John Hawks

I was reading The Scientist because RPM sent me to this article, titled "The Human Genome Project +5".

And yet the last five years, in Olson's view, have been "a period of a great grinding of gears, kind of shifting of gears." In the terms of the science historian Thomas Kuhn, it's been "a period of consolidation and more normal science." Others, such as Sydney Brenner of the Salk Institute, the Nobel Prize-winning pioneer of the worm, Caenorhabditis elegans, go further, worrying that the genome sequence and the growing lists of sequences and proteins and protein interactions and functional elements don't get very deep into such core problems of biology as the operations of the cell, of development from egg to adult, or the problem of consciousness. "We've become very geno-centric," says Brenner. "The cell must become the focus."

I would say this is pretty much correct -- there has been a long period of normal science in genetics lately, with new findings pretty much following one after another. There have been no revolutions coming out of the HGP.

But I think this scale of examination is a bit misleading. The HGP opened the deep end of the data pool, and we are still swimming in the toddler tank.

Consider what is happening in terms of new data:

One of the most dramatic efforts to push genomics into the realm of complex, multi-genic diseases is the five-year, $138 million haplotype map (HapMap) project, involving samples donated by Japanese, Han Chinese, Yoruba, and Americans of European descent. The project takes advantage of the fact that the millions of single nucleotide polymorphisms (SNPs) found in at least one percent of humans tend to pass between generations in blocks of DNA called haplotypes. The project announced its Phase 1 analysis in October 2005, and said that the analysis of Phase 2, already completed, would be published in 2006. Despite successes, such as using HapMap data to pinpoint a gene for macular degeneration, there remains controversy over HapMap's reach into domains such as rearrangements like deletions and reversals, or the numerous rare mutations that may be involved in diseases.

The minor variations are of central interest to Bentley of Solexa, who has specialized in rare variations. The HapMap, he says, has limitations, capturing only common variations in three target populations, missing the rare mutations. But it may provide a quick way to find more disease genes. Still, in three to five years, he says, the new sequencing machines should open the option of going after virtually all the many genes involved in a disease like diabetes. To be sure, the multiple sequences of patients and "controls" will have to square with what HapMap has found. "Everything that a HapMap captures should also be captured by a technology that aims to do better." Bentley, an early proponent, calls the HapMap "a real benchmark."

There seems to be a "Moore's law" for genome sequencing:

The workhorses of the 2001 human drafts have kept doubling their throughput about every 22 months over 15 years. In September, 454 reported that, in a single run, its system did a shotgun sequence and assembly of the microbe, Mycoplasma genitalium, in four hours. Claire Fraser's team at the Institute for Genomic Research took three months to work out Mycoplasma's sequence in 1995.

And there are gene expression microarrays and microRNA assays, as described in the article. For people who want to know about gene activity at every stage of life, in every type of cell, and in response to every external stimulus, the tools are in place to figure those things out.

As for myself, I think the accumulating data will have some revolutionary effects. These won't be in genetics itself -- I think the paradigms in place now in terms of gene interactions and regulation are very powerful. No doubt some new twists in gene sequence and function will be found, but I would guess that the current picture will expand rather than being overthrown.

But for other fields, I think genetics has some revolutionary power. Obviously genomic medicine has the potential to radically change the way we approach chronic conditions. And metagenomics is already changing the way biologists study microbial communities in all kinds of environments. It wouldn't surprise me if scientists working in places like the Foja Mountains work with DNA tag samples before they do traditional taxonomy on new species.

What will happen to anthropology as a result of the HapMap? There are surprises in store...

Peer review reviewed

Wed, 2006-02-08 23:18 -- John Hawks

There's an article about the problems with peer review by Alison McCook in The Scientist. I think it's a good summary of some of the difficulties with the review system with an exploding number of journal submissions, and it has many good quotes from journal editors.

Lawrence, based at the MRC Laboratory of Molecular Biology at Cambridge, UK, says his earlier papers were always published because he and his colleagues first submitted them to the journals they believed were most appropriate for the work. Now, because of the intense pressure to get into a handful of top journals, instead of sending less-than-groundbreaking work to second- or third-tier journals, more scientists are first sending their work to elite publications, where they often clearly don't belong.

Consequently, across the board, editors at top-tier journals say they are receiving more submissions every year, leading in many cases to more rejections, appeals, and complaints about the system overall. "We reject approximately 6,000 papers per year" before peer review, and submissions are steadily increasing, says Donald Kennedy, editor-in-chief of Science. "There's a lot of potential for complaints."

Since the interviews are mainly with big-time journal editors (Science, JAMA), they focus on the pressure to exaggerate findings and to burn papers from competitors. There's a great table that lists acceptance rates for some of these journals -- Science is less than 8 percent, PLoS Biology around 15 percent.

This was interesting to me:

Indeed, an abundance of data from a range of journals suggests peer review does little to improve papers. In one 1998 experiment designed to test what peer review uncovers, researchers intentionally introduced eight errors into a research paper. More than 200 reviewers identified an average of only two errors. That same year, a paper in the Annals of Emergency Medicine showed that reviewers couldn't spot two-thirds of the major errors in a fake manuscript. In July 2005, an article in JAMA showed that among recent clinical research articles published in major journals, 16% of the reports showing an intervention was effective were contradicted by later findings, suggesting reviewers may have missed major flaws.

A lot of the comments concerning signed reviews are that junior scientists will be afraid to write critical reviews of senior figures. There is some point to that, but there is no shortage of junior people willing to write letters when some senior person publishes an error, either.

And there are always blogs!

I don't really think that too many of these issues affect anthropological journals, although probably the press of submissions does at a slightly smaller scale. I do think that peer review would be improved by signing reviews, and I almost always give the editor the choice whether to keep mine anonymous. I hardly have to sign them, though; everybody knows they're twice as long as anybody else's!

Hybridization among Darwin's finches

Tue, 2006-02-07 22:57 -- John Hawks

This is an old paper by Peter and Rosemary Grant, from 2002:

Unpredictable Evolution in a 30-Year Study of Darwin's Finches

Peter R. Grant and B. Rosemary Grant

Evolution can be predicted in the short term from a knowledge of selection and inheritance. However, in the long term evolution is unpredictable because environments, which determine the directions and magnitudes of selection coefficients, fluctuate unpredictably. These two features of evolution, the predictable and unpredictable, are demonstrated in a study of two populations of Darwin's finches on the Galápagos island of Daphne Major. From 1972 to 2001, Geospiza fortis (medium ground finch) and Geospiza scandens (cactus finch) changed several times in body size and two beak traits. Natural selection occurred frequently in both species and varied from unidirectional to oscillating, episodic to gradual. Hybridization occurred repeatedly though rarely, resulting in elevated phenotypic variances in G. scandens and a change in beak shape. The phenotypic states of both species at the end of the 30-year study could not have been predicted at the beginning. Continuous, long-term studies are needed to detect and interpret rare but important events and nonuniform evolutionary change.

It seems like the purpose of the paper is to demonstrate the fluctuating evolution of the finches in response to environmental change. But I was drawn to it by the demonstration of biased gene flow from one species to the other:

The proportionally greater gene flow from G. fortis to G. scandens than vice versa has an ecological explanation. Adult sex ratios of G. scandens became male biased after 1983 (Fig. 4C) as a result of heavy mortality of the socially subordinate females. High mortality was caused by the decline of their principal dry-season food, Opuntia cactus seeds and flowers; rampantly growing vines smothered the bushes (16). G. fortis, more dependent on small seeds of several other plant species, retained a sex ratio close to 1:1 (Fig. 4C). Thus, when breeding resumed in 1987 after 2 years of drought, competition among females for mates was greater in G. fortis than in G. scandens. All 23 G. scandens females paired with G. scandens males, but two of 115 G. fortis females paired interspecifically. All their F1 offspring later bred with G. scandens (43) because choice of mates is largely determined by a sexual imprinting-like process on paternal song (42).

They conclude that this hybridization had an introgressive effect, acting in the same direction as selection for beak shape during the early 1990's, but continuing throughout the decade while there was little evidence for selection. There was little or no fitness loss to interspecific hybrids.

The case is distinctive because the effects of introgression on the characters of beak size and shape are consistently large enough to be measured each generation. This despite the fact that the proportion of F1 hybrids and first-generation backcrosses in the G. scandens population was never greater than 20 percent.

References:

Grant PR, Grant BR. 2002. Unpredictable evolution in a 30-year study of Darwin's finches. Science 296:707-711. Full text (subscription)

Evolving the electric swim

Tue, 2006-02-07 22:18 -- John Hawks

This is really cool:

The same genes that give sharks their sixth sense and allow them to detect electrical signals are also responsible for the development of head and facial features in humans, a new study suggests.

The finding supports the idea that the early sea creatures which eventually evolved into humans could also sense electricity before they emerged onto land.

The paper by Renata Freitas and colleagues is in Evolution and Development. Here is the suggestive part of the discussion:

Our finding that the rostral spread of the EphA4 expression domain prefigures the routes taken by mechanosensory and electrosensory axons is reminiscent of mouse ear innervation, in which EphA4 is expressed in the cells lining the auditory nerve pathway, where it directs axons to the cochlea (Pickles 2003). If this function is conserved in the shark laterosensory system, then EphA4 may be involved in guidance of sensory axons to electroreceptors and mechanoreceptors. Indeed, absence of EphA4 expression from ampullary placodes may also relate to the termination of growth cones at these positions. This may be important for both function and development of electroreceptors, as it has been suggested that the arrival of nerve fibers may induce formation of electroreceptive organs (although it is also possible that the placodes attract axons; Fritzsch et al. 1998).

Expression of EphA4 in the shark laterosensory system may represent a deeply conserved mechanism for establishing topographic maps of peripheral sensory inputs in vertebrates. In the mouse, EphA4 and EphrinA5 regulate development of the somatotopic map of projections from sensory whiskers to the barrel fields on the cortex (Vanderhaeghen et al. 2000). EphA4 has been shown to regulate thalamocortical projections, as well as the topographic projections of motor neurons from the spinal cord to the limb (Eberhart et al. 2000, 2002). Similar spatial patterning occurs in the auditory system, where topographic projections originating from the cochlea project to the nucleus magnocellularis, which in turn, innervates the nucleus laminaris in the brain to form a tonotopic map of high- to low-frequency sounds. Interestingly, EphA4 is expressed in a tonotopic gradient at the time when nucleus magnocellularis axons are forming synapses on the nucleus laminaris (Person et al. 2004). The association between expression of EphA4 and development of the shark electrosensory system suggests that EphA4 could play a role in establishing the topographic relationships between peripheral electroreceptors and their primary central targets. Regulation of EphA4 expression during development of the cephalic electrosensory system would therefore underlie how sharks localize the position of electrical stimuli relative to their spatial map of the body. This hypothesis is consistent with EphA4 playing a general role in the establishment of topographic maps during vertebrate embryogenesis (Vanderhaeghen et al. 2000; Yue et al. 2002; Dufour et al. 2003; Person et al. 2004).

This kind of deep genetic homology is not new -- one of the early insights of evo-devo was the widely shared genetic homologies underpinning sight in different lineages who "independently" evolved it. But this is interesting to me because electrosensory organs are easily seen as "exotic" characteristics of some lineages very distant to us. This study suggests that they may have served an important role in the early evolution of tetrapod ancestors. The genes that help generate electroreceptivity in sharks are still expressed in mammals, bent to other purposes.

The LiveScience article has an embryological note of caution from Glenn Northcutt, by the way:

Glenn Northcutt, a neuroscientist at the University of California at San Diego, who was not involved in the study, said the finding was interesting, but that more studies are needed before a direct link between neural crest cells and electroreceptors can be established.

"It still requires a definitive experiment, where the developing neural crest cells are marked with dye, the embryo develops and the dye clearly shows up in the electroreceptors," Northcutt said.

Evolution and Development is a really cool journal. Two of the other articles in the same issue relate to the sensory systems of cave fish -- a classic problem in evolution for which evo-devo approaches are giving interesting answers.

References:

Freitas R, Zhang GJ, Albert JS, Evans DH, Cohn MJ. 2006. Developmental origin of shark electrosensory organs. Evol Devel 8:74. DOI link

So which mythological paradise is it?

Tue, 2006-02-07 12:46 -- John Hawks

You've probably seen the story about the Foja Mountains in Papua New Guinea. Here are a couple of paragraphs:

JAKARTA, Indonesia - Describing it as the discovery of a "Lost World," conservation groups and Indonesia on Tuesday said an expedition to one of Asia's most isolated jungles had found several dozen new species of frogs, butterflies, flowers and birds.

"It's as close to the Garden of Eden as you're going to find on Earth," Bruce Beehler, a Conservation International scientist who led the expedition, said in a statement.

OK, so is it the "Lost World" or the "Garden of Eden"? Let's get this straight.

I say, it's the Garden of Eden if it has unicorns, and the Lost World if it has dinosaurs. Gretchen says it depends how bloodthirsty the animals are.

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Why are hybrids usually bad?

Mon, 2006-02-06 23:56 -- John Hawks

Two hypotheses, discussed by Burke and Arnold (2001):

The role of epistasis in adaptive evolution has been a controversial issue ever since Sewall Wright and R.A. Fisher first formalized their views in the early 1930s. According to Wright (113, 114), natural selection retains favorably interacting gene combinations. Therefore, as a result of the highly integrated nature of the genome, selection may lead to the production of what Dobzhansky (43) has termed "coadapted" gene complexes. In contrast, Fisher (48) argued that natural selection acts primarily on single genes, rather than on gene complexes. In Fisher's view, therefore, selection favors alleles that elevate fitness, on average, across all possible genetic backgrounds within a lineage. Such alleles have been termed "good mixers" (75). Regardless of the role of epistasis within lineages, however, negative epistasis in a hybrid genetic background, or hybrid incompatibility, is fully consistent with both the Wrightian and Fisherian worldviews. This is because allelic fixation occurs in any one lineage without regard to the compatibility (or lack thereof) of new alleles with those in any other lineage. Hybridization then produces a vast array of recombinant genotypes that have never before been subjected to selection. On average, these genotypes will be less well adapted than their parents, giving rise to some level of selection against hybrids.

Hybrid breakdown, or the reduction in fitness of segregating hybrid progeny that often results from intercrossing genetically divergent populations or taxa, has long been taken as evidence of unfavorable interactions between the genomes of the parental individuals (e.g., 39, 42, 43, 75, 80). The most widely accepted genetic model for the occurrence of such incompatibilities was first described by Bateson (15, as cited in 83), and later by Dobzhansky (39) and Muller (79, 80). In short, the Bateson-Dobzhansky-Muller (BDM) model assumes that an ancestral population consisting solely of individuals of the genotype aa/bb is broken into two parts that are temporarily isolated from each other. In one subpopulation, a new allele (A) is then assumed to arise at the first locus. Meanwhile, a new allele (B) is assumed to arise in the other subpopulation. Because individuals of the genotype aa/bb, Aa/bb, and AA/bb can interbreed freely, the A allele can then spread to fixation in the first subpopulation; likewise, individuals of the genotype aa/bb, aa/Bb, and aa/BB can interbreed freely, and the B allele spreads to fixation in the second subpopulation. However, although A is compatible with b, and B is compatible with a, the interaction of A with B is assumed to produce some sort of developmental or physiological breakdown, such that hybridization between the two subpopulations leads to the production of offspring with decreased levels of viability and/or fertility. Although this model focuses on negative interactions between differentiated regions of the nuclear genome, similar interactions between one or more regions of the nuclear genome and some component of the cytoplasm (e.g., the chloroplast or mitochondrial genome) could also play an important role in hybrid incompatibility. Unfortunately, the BDM model does not provide any mechanistic explanation as to how mutations that are neutral (or beneficial) within a given lineage will produce strongly disadvantageous incompatibilities when combined in a hybrid background (Burke and Arnold 2001, emphasis added).

References:

Burke JM, Arnold ML. 2001. Genetics and the fitness of hybrids. Annu Rev Genet 35:31-52. DOI link

New French Paleolithic cave art

Mon, 2006-02-06 22:06 -- John Hawks

The AP is reporting on a new cave art find in France.

PARIS - Cave drawings thought to be older than those in the famed caves of Lascaux have been discovered in a grotto in western France, officials from the Charente region said Sunday.

A first analysis by officials from the office of cultural affairs suggests the drawings were made some 25,000 years ago, Henri de Marcellus, mayor of the town of Vilhonneur where the cave is located, told France-Info radio.

No details or pictures. The BBC has some quotes from the discoverer:

Gerard Jourdy, 63, said he found human and animal remains in the chamber in the Vilhonneur forest, in caves once used to dispose of animal carcasses.

The paintings included a hand in cobalt blue, he told AFP news agency.

The discovery was made in November, but kept secret while initial examinations were carried out.

Mr Jourdy also said he saw a sculpture of a face made from a stalactite - which would be a scientific first for the era, but experts were dubious about this claim, AFP says.

"In a small chamber I found the bones of two hyenas - complete skeletons, which is rare. And I saw human bones amid the debris - tibias, vertebrae and shoulder-blades," he told the news agency.

"Then in the bigger chamber there was this hand - very beautiful, very delicate. There was just the one in cobalt blue. When you come into the chamber it is like it is greeting you. It's incredible."

I wonder if those bones are really human. I guess we'll find out.

UPDATE (2/6/06): The New York Times has a short item including this:

Michel Bilaud, the governor of the department of Charente, expressed doubt about the art. "There are traces of human occupation," he said. "There are bones, and there are lines on the wall. There is a print of a hand. But for the rest, it is just marks. There is nothing figurative."

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"Africa": the game

Mon, 2006-02-06 21:44 -- John Hawks

MTV News has an article describing a new massively multiplayer online game, called "Africa". The game will be set in "a land of 13th century African civilization and mythology", encompassing the entire continent.

"We felt very strongly that video games can help increase understanding and education about Africa and get the unmotivated public fired up about what is going on with Africa," said John Sarpong, grandson of Ashanti king Prampeh of Ghana, exiled by the British from 1895 to 1924. Sarpong has spent recent years running Africast, a company that broadcasts African programming over the Internet.

Less reserved, Adam Ghetti, the teenage creative director at Rapid Reality, the company actually creating the game, said he hopes the game will right some wrongs. "The white American board developers of the large MMO development companies out there right now don't honestly have the right background and knowledge on the continent of Africa and its lore, mythology and rich history, and quite honestly neither did I," said Ghetti, who is white. "They just don't teach it over here." The game is designed, in part, to change that.

It sounds a bit more involved than "The Oregon Trail".

I'm really attracted to this concept of immersion into a historic world. Of course, the quality of the experience is only as good as the quality of the AI interactors, and the other players.

Now, I suppose some people will get into the experience like Civil War reenactors, but for the most part, they won't share culture with 13th century Africans. More and more, people are experiencing these pseudo-ethnographic simulations; I wonder where they are going to end up:

"Africa" will be a vessel for Ghetti and Spaight's ambition. They want a virtual world that functions dynamically: antelope that find new pastures when grassland is scorched, drum music fully customizable by players and used -- as in real ancient Africa -- as an alternate langue. They want players to be able to become famous and change the map. More flexible than the mostly developer-controlled "World of Warcraft" but more restrained than the free-living "Second Life," "Africa" would let a tribe of 100 players establish their own officially recognized empire, but only after the equivalent of 12 hours of play across 48 weeks. For the less hard-core, play can be done casually as a fighter, merchant, musician or even a human who can turn into a bird.

Ghetti said he thinks the setting will present gamers a welcome and surprisingly rich change of scenery. "The African mythology back from 1200 to 1400 A.D. is thousands of times richer than the J.R.R. Tolkien series of novels," he said. "Don't get me wrong, he was an amazing individual with brilliant ideas. But that's been milked for 80 years now."

Yes, it will be much more useful to have people learning about the actual landscape of historic Africa than the fictional landscape of Middle Earth. It should be interesting -- MMO games are increasingly involving real-life trade in currency and skills. The game creators say they hope the game will be played in Africa. You can be pretty sure it will be played in China -- a lot, since Chinese companies employ game players to get virtual goods to be sold on EBay.

Imagine the 45-year-old white stocktrader paying 30 seventeen-year-old Chinese kids to help him become the next king of the Ashanti.

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Mirror neurons in video gaming

Mon, 2006-02-06 13:15 -- John Hawks

For those interested in the sociological side of neuroscience, GameSpot has a quick review of mirror neurons and their relevance to video games.

Neuroscience and video games. What do they have to do with each other? Aside from whatever research went into crafting games like Psi-Ops and Psychonauts, it doesn't seem like the two subjects have much in common. Sure, neuroscience is the study of the brain, and despite what everyone tells you, you do use your brain when you're playing video games. But what are the chances that the latest neuroscientific research is going to be of any interest to the game industry? Well, if you've been following the (relatively) recent work on mirror neurons, then you would realize that neuroscience is about to have a huge impact--if not on video games, then on the discussions we have about them--for a long time to come.

The emphasis on the article is the possible long-term effects of violent video games on violent crime rates. To that end, it discusses the possible effects of mirror neurons in familiarizing people with experiences that they do not themselves actually do, as well as "super mirror" neurons that may regulate those perceptions.

We can tell if someone is watching a television by the way that person is facing it--even if we can't see or hear if the television is even on. It also means that we can experience the mental states associated with actions without ever having to perform those actions. In video games, in particular, it's like we're automatically empathizing with what is happening on the screen as if we were the video game characters ourselves. If you've ever had a particularly heart-palpitating race in Burnout, surely you can relate.

"We can tell if someone is watching a television..." certainly gives a hint about the evolutionary history of the mirror neurons, but they are strange beasts.

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Hybrid swarms

Mon, 2006-02-06 11:17 -- John Hawks

I found this paper by E. B. Taylor and colleagues from a link on evolgen. The paper is titled "Speciation in reverse", and it is about the loss of distinction between two stickleback morphs in Enos Lake, British Columbia.

From the abstract:

Bayesian analyses of population structure in a sample collected in 1994 indicated two genetically distinct populations in Enos Lake, but only a single genetic population was evident in 1997, 2000, and 2002. In addition, genetic analyses of samples collected in 1997, 2000, and 2002 showed strong signals of 'hybrids'; they were genetically intermediate to parental genotypes. Our results support the idea that the Enos Lake species pair is collapsing into a hybrid swarm.

I love this paragraph from the introduction:

The persistence of sympatric species that still occasionally hybridize implies that a dynamic balance exists between the occasional production of hybrids and their removal by natural selection. Presumably, there have been cases where completion of speciation in sympatry fails because gene flow overwhelms factors promoting divergence. Perhaps more infrequently, speciation proceeds to completion only to be undone later when environmental conditions change. Such reversals can occur, for instance, if the fitness of hybrids is suddenly improved under the new environmental conditions (e.g. Grant and Grant 2002). Alternatively, environmental cues upon which premating isolation is based may suddenly be altered, leading to a burst of hybridization that selection can no longer overcome (discussed in Coyne and Orr 2004). In either instance, the frail integrity of species that lack complete postzygotic isolation demonstrates the contribution of the environment to their maintenance and, perhaps, provides insight to the identity of factors that initiate speciation.

"The frail integrity of species that lack complete postzygotic isolation."

There is probably a reason why these particular populations have merged. The authors suspect that the introduction of nonnative crayfish may be a factor -- the crayfish eat some of the deep-water sticklebacks and may compete with them for food.

These stickleback populations have postglacial origins -- the age of Enos lake. Elsewhere, the divergence of stickleback populations in postglacial lakes has been found to be independently repeated, in terms of the origin of distinct benthic and limnetic (deep and surface) forms.

The most interesting aspect of this for me is the initial morphological divergence of the populations. They clearly have been adapting to distinct roles in these lakes even with substantial possibility of ongoing gene flow -- even if they were partially isolated by their use of different breeding strategies and water depths.

Highly adaptable. Similar morphological forms repeatedly evolve in similar habitats. Rapid speciation with ongoing genetic exchanges. Rapid collapse of morphological diversity (and genetic distinctiveness) in response to environmental change.

References:

Taylor EB, Boughman JW, Groenenboom M, Sniatynski M, Schluter D, Gow JL. 2006. Speciation in reverse: morphological and genetic evidence of the collapse of a three-spined stickleback (Gasterosteus aculeatus) species pair. Mol Ecol 15:343. DOI link

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Rediscovering the obvious

Sun, 2006-02-05 20:00 -- John Hawks

I was just discussing with someone today the merits of taking an old idea and giving it a new name, and now I find out from the NY Times just what a good strategy it can be:

It may seem odd that scientists in the Internet age spend years on a line of research, even bet their careers on it, without having first determined that their mountain had not already been climbed. But Dr. Stigler said that scientists often are ignorant of the work being done by others in their field, and searches of scientific literature can be hard to conduct. Web search engines, for example, look for words, not ideas, and Dr. Vohra said he discovered that every researcher who had made his discovery had given it a different name and description.

In 1957, for example, a statistician named James Hanna called his theorem Bayesian Regret. He had been preceded by David Blackwell, also a statistician, who called his theorem Controlled Random Walks. Other, later papers had titles like "On Pseudo Games," "How to Play an Unknown Game," "Universal Coding" and "Universal Portfolios," Dr. Vohra said, adding, "It's not obvious how you do a literature search for this result."

The idea in question is that random stock buyers can outperform professionals with a simple buy-and-hold strategy. I especially like "Bayesian Regret"; it looks like somebody's dissertation title.

There's always this;

For example, there is the oft-told story about Larry Shepp, a famous mathematician at Rutgers University. Dr. Shepp, when told that a piece of work he thought was his discovery actually duplicated another mathematician's breakthrough, replied: "Yes, but when I discovered it, it stayed discovered."

The Templeton review

Fri, 2006-02-03 11:50 -- John Hawks

The Yearbook of Physical Anthropology has a new review of the genetic evidence for modern human origins by Alan Templeton. The paper is 27 journal pages, and they are full of detail -- especially after the section describing basic coalescent theory.

I'll be going through this paper in the next few days and highlighting some of the issues it raises. In the meantime, here are some quotes from the Washington University press release:

"The 'Out of Africa' replacement theory has always been a big controversy," Templeton said. "I set up a null hypothesis and the program rejected that hypothesis using the new data with a probability level of 10 to the minus 17th. In science, you don't get any more conclusive than that. It says that the hypothesis of no interbreeding is so grossly incompatible with the data, that you can reject it."

...

The new data confirm an expansion out of Africa to 700,000 years ago that was detected in the 2002 analysis.

"Both (the 1.9 million and 700,000 year) expansions coincide with recent paleoclimatic data that indicate periods of very high rainfall in eastern Africa, making what is now the Sahara Desert a savannah," Templeton said. "That makes the timing very amenable for movements of large populations through the area."

Found via Dienekes, who seems to be one step ahead of me this week!

References:

Templeton AR. 2005. Haplotype trees and modern human origins. Yrbk Phys Anthropol 128(S41):33-59. DOI link

"'Forever' is not a mathematically tractable quantity"

Fri, 2006-02-03 10:17 -- John Hawks

That's my favorite line from this article in the Economist about Bayesian logic.

From Wikipedia, a short introduction to Bayesian logic:

For example, Laplace estimated the mass of Saturn using Bayesian methods. However, on the frequency interpretation of probability the laws of probability cannot be applied to this problem. This is because the mass of Saturn isn't a well defined random experiment or sample. From what population is the mass of Saturn taken? In what sense is Saturn picked at random from that population? Similarly, when comparing two hypotheses and using the same information, frequency methods would typically result in the rejection or non-rejection of the original hypothesis with a particular degree of confidence, while Bayesian methods would yield statements that one hypothesis was more probable than the other or that the expected loss associated with one was less than the expected loss of the other.

The Economist article is a good short introduction to what Bayesian logic is, and it follows some interesting research that indicates that human minds are especially good at it:

Dr Griffiths and Dr Tenenbaum conducted their experiment by giving individual nuggets of information to each of the participants in their study (of which they had, in an ironically frequentist way of doing things, a total of 350), and asking them to draw a general conclusion. For example, many of the participants were told the amount of money that a film had supposedly earned since its release, and asked to estimate what its total "gross" would be, even though they were not told for how long it had been on release so far.

Besides the returns on films, the participants were asked about things as diverse as the number of lines in a poem (given how far into the poem a single line is), the time it takes to bake a cake (given how long it has already been in the oven), and the total length of the term that would be served by an American congressman (given how long he has already been in the House of Representatives). All of these things have well-established probability distributions, and all of them, together with three other items on the list -- an individual's lifespan given his current age, the run-time of a film, and the amount of time spent on hold in a telephone queuing system -- were predicted accurately by the participants from lone pieces of data.

By "accurately", they appear to mean that the prior distribution was reconstructable from the responses of the 350 people, not that every person guessed the right answer.

The "forever" line comes from this passage:

There were only two exceptions, and both proved the general rule, though in different ways. Some 52% of people predicted that a marriage would last forever when told how long it had already lasted. As the authors report, "this accurately reflects the proportion of marriages that end in divorce", so the participants had clearly got the right idea. But they had got the detail wrong. Even the best marriages do not last forever. Somebody dies. And "forever" is not a mathematically tractable quantity, so Dr Griffiths and Dr Tenenbaum abandoned their analysis of this set of data.

Personally, I think it's romantic that 52 percent of people predicted that marriage would last forever!

I suppose that helps to underlie the last two paragraphs:

How the priors are themselves constructed in the mind has yet to be investigated in detail. Obviously they are learned by experience, but the exact process is not properly understood. Indeed, some people suspect that the parsimony of Bayesian reasoning leads occasionally to it going spectacularly awry, with whatever process it is that forms the priors getting further and further off-track rather than converging on the correct distribution.

That might explain the emergence of superstitious behaviour, with an accidental correlation or two being misinterpreted by the brain as causal. A frequentist way of doing things would reduce the risk of that happening. But by the time the frequentist had enough data to draw a conclusion, he might already be dead.

Which may help to explain why it takes a certain size and structure of population to develop science -- probably including writing, so it's harder to ignore information that someone else collects.

Anyway, read the article to find out the problem people had guessing the length of the reigns of Egyptian Pharaohs.

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More on myosin mutations

Wed, 2006-02-01 22:53 -- John Hawks

Melanie McCollum and colleagues have a short paper in JHE about the evolution of MYH16, the myosin gene associated with masticatory musculature.

Much of the article is devoted to debunking the connection of MYH16 to the expansion of the brain. There's not much to that story -- muscles beating down on the skull don't inhibit brain growth. And they discuss the implications of the revised date estimate for the deactivation of the gene by Perry and colleagues (2005).

But by far the most interesting part of this is their discussion of the effects of the evolution of masticatory myosin in other mammals:

With respect to interspecific variation in the expression of "masticatory" myosin, it is important to note that humans are not unique in their failure to express this particular isoform. "Masticatory" myosin is lacking in the jaw-closing muscles of a number of mammals, most notably ungulates, rodents, rabbits, and kangaroos (Kang et al., 1994, Sfondrini et al., 1996, Hoh, 2002 and Qin et al., 2002). Comparative genetic studies suggest that the masticatory MyHC gene originated through duplication of an ancestral striated MyHC gene expressed in the mandibular arch musculature of early gnathostomes, and that it has since been retained as the primitive phenotype in vertebrates (Qin et al., 2002). Functional loss of masticatory myosin in a number of non-carnivorous mammalian species is believed to have followed shifts in dietary strategies that ultimately freed these taxa from the need for powerful jaw closure. As a consequence, these taxa are believed to have replaced their masticatory myosin with functionally more appropriate myosin isoforms (e.g., slow/beta-cardiac fibers in ungulates, Kang et al., 1994; fast MyHCs in rodents, Sfondrini et al., 1996) (McCollum et al. 2006, references in original).

This would seem to support the hypothesis that changing masticatory function in hominids (caused by dietary changes) favored the gene's deactivation. But deactivation might have preceded the conversion of the gene into a pseudogene:

However, the very fact that muscle fibers readily change their myosin heavy chain expression suggests that masticatory myosin in hominids could very well have been significantly, if not totally downregulated prior to its conversion to a pseudogene. If this were the case, inactivation of the MYH16 gene would have had little impact on the muscles of mastication of early hominids and far less severe consequences for its carriers. In fact, the introduction of a nonsense mutation in the MYH16 gene may have been below the threshold of selection. If this alternative is correct, then the real question of interest is whether the change in masticatory function that occurred during hominid evolution and that led to MYH16 downregulation and inactivation was diet-related, as has been recently suggested (Hoh, 2002), or instead reflected changes in social behaviors that would have eliminated the need for an aggressive bite, as was suggested over 20 years ago (Rowlerson et al., 1983).

OK, so we have first the question of the date (2.4 million vs. 5.3 million years ago), and second the question of whether the functional change -- whatever it was -- preceded the date. So this could reflect anything from change associated with hominid origins up to the origin of Homo.

McCollum et al. make clear in their description of muscle fiber function that we may be looking not at a muscle decrease but a functional change. In which case, the deactivation might have accompanied greater masticatory force instead of less.

What a mess.

References:

McCollum MA, Sherwood CC, Vinyard CJ, Lovejoy CO, Schachat F. 2006. Of muscle-bound crania and human brain evolution: The story behind the MYH16 headlines. J Hum Evol (in press) DOI link

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Mojokerto site rediscovered?

Wed, 2006-02-01 22:29 -- John Hawks

An upcoming paper in Journal of Human Evolution by O. F. Huffman and colleagues reports on a possible location for the Mojokerto skull. A 1994 paper by Carl Swisher and colleagues dated rock from the supposed site to 1.81 million years ago.

This paper finds that the real site is a bit above that dated horizon. The abstract:

The fossil calvaria known as the Mojokerto child's skull was discovered in 1936, but uncertainties have persisted about its paleoenvironmental context and geological age because of difficulties in relocating the discovery site. Past relocation efforts were hindered by inaccuracies in old base maps, intensive post-1930s agricultural terracing, and new tree and brush growth. Fortunately geologic cross sections and site photographs from 1936-1938 -- not fully utilized in past relocation fieldwork -- closely circumscribe site geography and geology. These documents match the conditions at just one sandstone outcrop. It is situated on the southern margin of a topographic nose at the upper end of a 18 m-wide gully (0663760 m E, 9183430 m N, UTM Zone 49M), 15 m southeast of the Kumai et al. (1985) relocation. The relocated discovery bed is 3.3 m of fossiliferous pebbly sandstone, a river-channel deposit cut into tuffaceous mudstone. The sandstone and mudstone beds correspond to original site descriptions. Pebbly sandstone is also found within the skull.

The calvaria is well-preserved and taphonomically similar to large and fragile specimens found among several hundred vertebrate fossils excavated from the sandstone in 2001-2002. Since no well-preserved fossils were found intact at the surface of the sandstone, the good condition of the Mojokerto skull suggests that it was buried fully when discovered. The relocated hominin bed is the uppermost fluvial sandstone of a marine-deltaic sequence in the upper Pucangan Formation. The Mojokerto child probably died along the ancient seacoast, judging from the large extent of the deltaic facies and evidence that the calvaria experienced minimal transport. The relocated discovery bed is 20 m stratigraphically above the horizon from which the widely cited 1.81 +- 0.04 Ma 40Ar39Ar date for the skull (Swisher et al., 1994, Science 263, 1118) was obtained. Additional field and laboratory results will be required to determine the skull's age.

The paper gives a good history of attempts to find the original excavation site. An interesting heterogeneity of the matrix fill inside the skull (assessed by CT) also factors in the story.

After a long discussion of the complexities in dating the site, they conclude:

In summary, additional field and analytical results are needed to date the Mojokerto fossil more exactly than latest Pliocene or early-mid Pleistocene in age. The 0.3 Ma difference between the 40Ar/39Ar and fission-track age determinations must be resolved. For any of these radioisotopic dates to be considered other than a maximum age, better evidence must be advanced to show that the dated material was erupted shortly before deposition at Perning. Additional paleontological and magnetostratigraphic control and radioisotopic dating would seem to be required. Geochronological conclusions have to be evaluated further in terms of the potential for temporal stratigraphic breaks in the section, rates of deposition, and the regional stratigraphic (including sequence stratigraphic) context.

I don't suppose there will ever be a very good date for the specimen. But it's impressive the amount of work there has been on it in the past several years.

References:

Huffman OF, Zaim Y, Kappelman J, Ruez DR Jr, de Vos J, Rizal Y, Aziz F, Hertler C. 2006. Relocation of the 1936 Mojokerto skull discovery site near Perning, East Java. J Hum Evol (in press). DOI

Swisher CC 3rd, Curtis GH, Jacob T, Getty AG, Suprijo A, Widiasmoro. 1994. Age of the earliest known hominids in Java, Indonesia. Science 263:1118-1121. PubMed

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Drifting away from selection

Wed, 2006-02-01 21:42 -- John Hawks

Following up on yesterday's post on annoying misconceptions, I noticed Razib had posted his own candidate:

My problem is not an misconception, it is a pet peeve. As I've noted before, random genetic drift is a catchall explanation for everything.

Well, I thought that was worth a post of its own instead of an update, because it probably annoys me even more than the species divergence thing.

It is a periodic revelation for me how many committed Darwinists don't use or understand natural selection. I think they place natural selection somewhere between Santa Claus and the Tooth Fairy -- useful to explain a few really strange phenomena, but pretty much irrelevant to evolution.

There is, of course, some reason to be cautious about selection. A little selection goes a long way toward explaining almost any pattern of evolution. Selection can make populations stay the same, and it can make them change. It can make them change fast, or it can make them change slowly. And at the genetic level, there are good reasons to suppose that many nucleotide changes don't have a phenotypic effect -- necessary for them to be selected. So it is reasonable a lot of the time to take neutrality (and therefore, genetic drift) as a null hypothesis for change.

Null hypotheses aren't there to be believed, they are there to be tested! Neutrality is a better null hypothesis because the hypothesis of some kind of selection is harder to refute. But neutrality is pretty hard to refute too, at least for the kind of evidence we usually have at hand.

It doesn't help at the molecular level that clearly non-neutral patterns of variation can be explained by extreme demographic changes (and no selection), or selection. Which hypothesis do we choose then? Most people pick neutrality, but not always for good reasons.

Nor does it help that morphological change over long time spans tends to average to very small amounts per unit time. That pattern of change is thoroughly consistent with genetic drift, but equally consistent with slowly changing stabilizing selection. And depending on the density of fossil sampling, it is often consistent with occasional pulses of strong directional selection.

Testing the difference between these hypotheses statistically is murderously hard with fossil samples. So what do we accept provisionally?

Myself, I'm a natural selection man. Accident is overrated.

You know, there's something depressing about collecting a bunch of annoying misconceptions -- and I suppose reading all of them must be sort of annoying itself. Of course, there's always the hope that writing about them might have some effect ...

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Neandertals

For years, I've worked on their bones. Now I'm working on their genes. Read more about the science studying these ancient people.

Denisova

From a finger bone of an ancient human came the record of a completely unexpected population. My lab is working on the science of the Denisova genome.

Acceleration

The advent of agriculture caused natural selection to speed up greatly in humans. We're uncovering some of the ways that populations have rapidly changed during the last 10,000 years.

Malapa

Just outside Johannesburg, the Malapa site is producing some of the most exciting finds in human evolution. This site is the headquarters of the Malapa Soft Tissue Project.