Molecular dating when rates vary

Molecular-dating techniques potentially enable us to estimate the time of origin of any biological lineage. Such techniques were originally premised on the assumption of a 'molecular clock'; that is, the assumption that genetic change accumulated steadily over time. However, it is becoming increasingly clear that constant rates of molecular evolution might be the exception rather than the rule. Recently, new methods have appeared that enable the incorporation of variable rates into molecular dating. Direct comparisons between these methods are difficult, because they differ in so many respects. However, the assumptions about rate change on which they rely fall into a few broad categories. Improving our understanding of molecular evolution will be an important next step towards evaluating and improving these methods.     

Molecular dating in the genomic era

The comparison of DNA and protein sequences of extant species might be informative for reconstructing the chronology of evolutionary events on Earth. A phylogenetic tree inferred from molecular data directly depicts the evolutionary affinities of species and indirectly allows estimating the age of their origin and diversification. Molecular dating is achieved by assuming the molecular clock hypothesis, i.e., that the rate of change of nucleotide and amino acid sequences is on average constant over geological time. If paleontological calibrations are available, then absolute divergence times of species can be estimated. However, three major difficulties potentially hamper molecular dating : (1) a limited sample of genes and organisms, (2) a limited number of fossil references, and (3) pervasive variations of molecular evolutionary rates among genomes and species. To circumvent these problems, different solutions have been recently proposed. Larger data sets are built with more genes and more species sampled through the mining of an increasing number of genomes. Moreover, independent key fossils are identified to calibrate molecular clocks, and the uncertainty on their age is integrated in subsequent analyses. Finally, models of molecular rate variations are constructed, and incorporated in the so-called relaxed molecular clock approaches. As an illustration of these improvements, we mention that the debated age of the animal (bilaterian metazoans) diversification may have occurred between 642-761 million years ago (Mya), roughly 100 Ma before the Cambrian explosion. Among mammals, the initial diversification of major placental groups may have taken place around 100 Mya, well before the Cretaceous/Tertiary boundary marking the extinction of dinosaurs.     

Molecular dating and biogeography of fig-pollinating wasps

Figs and fig-pollinating wasps are obligate mutualists that have coevolved for over 60 million years. But when and where did pollinating fig wasps (Agaonidae) originate? Some studies suggest that agaonids arose in the Late Cretaceous and the current distribution of fig-wasp faunas can be explained by the break-up of the Gondwanan landmass. However, recent molecular-dating studies suggest divergence time estimates that are inconsistent with the Gondwanan vicariance hypothesis and imply that long distance oceanic dispersal could have been an important process for explaining the current distribution of both figs and fig wasps. Here, we use a combination of phylogenetic and biogeographical data to infer the age, the major period of diversification, and the geographic origin of pollinating fig wasps. Age estimates ranged widely depending on the molecular-dating method used and even when using the same method but with slightly different constraints, making it difficult to assess with certainty a Gondwanan origin of agaonids. The reconstruction of ancestral areas suggests that the most recent common ancestor of all extant fig-pollinating wasps was most likely Asian, although a southern Gondwana origin cannot be rejected. Our analysis also suggests that dispersal has played a more important role in the development of the fig-wasp biota than previously assumed.     

Primatology: advanced ape technology

New findings from African rainforests show chimpanzees to have impressively advanced technology. They make tools of vegetation to harvest termites as in East and West Africa, but some apes in Central Africa show different techniques and tool sets geared for different tasks.     

Paleoanthropological applications of amino acid racemization dating of fossil bones and teeth

The general principals of the amino acid racemization based dating technique are discussed. The results obtained at Olduvai Gorge (Tanzania) and Zhoukoudian (China) are presented as an illustration of the paleoanthropological application of racemization dating. Tooth enamel provides the best material for the racemization dating of Middle to Lower Pleistocene deposits, although in some cases bones also yield reliable ages. The racemization method provides a means of dating teeth and bones which are too old for radiocarbon dating. Only small samples are required and processing procedures are straightforward. The technique is particularly useful in paleoanthropological studies since hominid fossils can be directly dated.     

Electron spin resonance dating of human bones from Brazilian shell-mounds (Sambaquís)

From the fingertip pattern analysis of 125 families with 376 children from Velanadu Brahmin and 100 families with 286 children from Telaganya Brahmin caste sects of Andhra Pradesh, India, the occurrence of an arch on at least one of the ten finger tips is shown to be aggregated in families. The mode of aggregation is in conformity with a major dominant gene with 0.28 chance of penetrance. This is to some extent contradictory to the result published from the analysis of a single Habbanite Jewish pedigree. Some implications of this discordant result are also discussed.     

Molecular dating and biogeography of the early placental mammal radiation

The timing and phylogenetic hierarchy of early placental mammal divergences was determined based on combined DNA sequence analysis of 18 gene segments (9779 bp) from 64 species. Using rooted and unrooted phylogenies derived from distinct theoretical approaches, strong support for the divergence of four principal clades of eutherian mammals was achieved. Minimum divergence dates of the earliest nodes in the placental mammal phylogeny were estimated with a quartet-based maximum-likelihood method that accommodates rate variation among lineages using conservative fossil calibrations from nine different nodes in the eutherian tree. These minimum estimates resolve the earliest placental mammal divergence nodes at periods between 64 and 104 million years ago, in essentially every case predating the Cretaceous-Tertiary (K-T) boundary. The pattern and timing of these divergences allow a geographic interpretation of the primary branching events in eutherian history, likely originating in the southern supercontinent Gondwanaland coincident with its breakup into Africa and South America 95-105 million years ago. We propose an integrated genomic, paleontological, and biogeographic hypothesis to account for these earliest splits on the placental mammal family tree and address current discrepancies between fossil and molecular evidence.     

Molecular phylogenetic and molecular dating of the New Zealand Gleicheniaceae

The Gleicheniaceae are an ancient family of ferns, with three of the six extant genera occurring in New Zealand:Dicranopteris, Gleichenia, andSticherus. The biogeographic origins of this family in New Zeland are unknown, and the taxonomy ofGleichenia in particular is still unclear. To address aspects of these two issues, DNA sequences from thetrnL-trnF locus and therbcL gene were produced for all of the common Gleicheniaceae species in New Zealand, as well as forGleichenia alpina from Tasmania andSticherus cryptocarpus from Chile. SeveraltrnL-trnF haplotypes were found amongst New ZealandG. dicarpa. One of these haplotypes was also observed in TasmanianG. alpina, while the other New ZealandG. dicarpa trnL-trnF haplotypes were more similar to those ofG. microphylla. These results suggest the taxonomy of New ZealandGleichenia may be more complex than presently recognized. Molecular dating of therbcL sequences with the program r8s rejected vicariant explanations for the disjunct distributions between New Zealand and elsewhere for each ofDicranopteris, Gleichenia, andSticherus. However, the direction of the inferred long-distance dispersal was not resolved.     

Molecular charcteristics of chicken kidney arginase.

Chicken kidney contains two arginases with different sedimentation coefficients and substrate specificity. The ligher of these arginases, which hydrolyses only L-arginine, has been purified about 3000-fold. Like the "ureotelic" arginase, developed in chicken liver after starvation, it displays many of the properties of the arginase of the "ureotelic" species. This seems to exclude the possibility that ureotelism and uricotelism are characterized by a specific type of arginases. Both liver and kidney arginases are located in the mitochondrial matrix. The rate of hydrolysis of arginine thus not only depends on the arginase activity but also on the rate of transport of arginine into the matrix. This last process therefore is of regulatory significance.     

Molecular analysis of chicken immune response genes

We have recently isolated immune response genes of the major histocompatibility B complex of the chicken (the B-L beta genes) by cross-hybridization in low stringency with an HLA class II beta chain probe. After reviewing the main results obtained, we present a detailed analysis of the region flanking the first gene characterized, B-L beta III. By Southern blot analysis with exon-specific probes, we demonstrate the presence of another related B-L beta gene 10 kb on the 3' side of B-L beta III, the B-L beta V gene. Moreover, retrospective analysis of the phage clones initially isolated with the HLA-DQ beta probe, using a chicken class I probe that we isolated by chromosome walking from the B-L beta genes, indicates that the B-L beta III gene is closely linked on its 5' side to a class I gene, B-FVI.     

Molecular cloning of chicken prepro-orexin cDNA and preferential expression in the chicken hypothalamus

Chicken prepro-orexin cDNA has been cloned, sequenced and characterized. The predicted amino acid sequence of chicken prepro-orexin cDNA revealed that orexin-A and -B are highly conserved among vertebrate species. In situ hybridization and immunohistochemistry localized orexin-positive cell bodies in the periventricular hypothalamic nucleus extending into the lateral hypothalamic area. Comparisons of orexin gene expression in the brains of 24-h-fasted and ad libitum-fed chickens were made using semi-quantitative RT-PCR. No significant differences in orexin mRNA expression were observed.     

Molecular phylogenetic dating of asterid flowering plants shows early Cretaceous diversification

We present a phylogenetic dating of asterids, based on a 111-taxon tree representing all major groups and orders and 83 of the 102 families of asterids, with an underlying data set comprising six chloroplast DNA markers totaling 9914 positions. Phylogenetic dating was done with semiparametric rate smoothing by penalized likelihood. Confidence intervals were calculated by bootstrapping. Six reference fossils were used for calibration. To explore the effects of various sources of error, we repeated the analyses with alternative dating methods (nonparametric rate smoothing and the Langley-Fitch clock-based method), alternative tree topologies, reduced taxon sampling (22 of the 111 taxa deleted), partitioning the data into three genes and three noncoding regions, and calibrating with single reference fossils. The analyses with alternative topologies, reduced taxon sampling, and coding versus noncoding sequences all yielded small or in some cases no deviations. The choice of method influenced the age estimates of a few nodes considerably. Calibration with reference fossils is a critical issue, and use of single reference fossils yielded different results depending on the fossil. The bootstrap confidence intervals were generally small. Our results show that asterids and their major subgroups euasterids, campanulids, and lamiids diversified during the Early Cretaceous. Cornales, Ericales, and Aquifoliales also have crown node ages from the Early Cretaceous. Dipsacales and Solanales are from the Mid-Cretaceous, the other orders of core campanulids and core lamiids from the Late Cretaceous. The considerable diversity exhibited by asterids almost from their first appearance in the fossil record also supports an origin and first phase of diversification in the Early Cretaceous.