Growth and foraging consequences of facultative paedomorphosis in the tiger salamander,Ambystoma tigrinum nebulosum

Summary Facultative paedomorphosis in salamanders occurs when larvae respond to varying environmental conditions by either metamorphosing into terrestrial metamorphic adults or retaining their larval morphology to become sexually mature paedomorphic adults. Several hypotheses have been proposed for the evolutionary maintenance of this environmentally induced dimorphism, but few data are available to assess them adequately. We studied a montane population of the tiger salamander,Ambystoma tigrinum nebulosum, and measured the adult growth rate and body condition across three growing seasons to assess the relative costs and benefits of each morph. Metamorphic adults grew more than paedomorphic adults in terms of snout—vent length across years and in weight within years. Dietary analyses and foraging experiments revealed some of the proximate factors that may underlie these differential growth patterns. Across all prey, metamorphs had significantly higher biomass and calories per stomach sample than paedomorphs. Metamorphic diets primarily consisted of the fairy shrimpBranchinecta coloradensis, whereas paedomorphic diets contained a variety of benthic and terrestrial invertebrates. Foraging experiments revealed that both morphs are more successful at capturing fairy shrimp relative to other prey types and both show high electivity toward this prey. However, fairy shrimp occurred only in non-permanent ponds and thus are inaccessible to paedomorphs, which can survive only in permanent ponds. Paedomorphs also experience higher levels of intraspecific competition with large larvae in permanent ponds than metamorphs do in non-permanent ponds. Thus, metamorphs obtain a growth advantage over paedomorphs by foraging in non-permanent ponds that contain fairy shrimp and have reduced intraspecific competition. These results suggest that paedomorphs should have decreased fitness relative to metamorphs, primarily because metamorphs can move into the best habitats for growth. The net fitness effect of morph-specific differences in dispersal depend on whether there are trade-offs with other life history traits. Nonetheless, because the relative benefit of metamorph dispersal will change with environmental conditions in permanent ponds and the surrounding habitat, the relative fitness payoff to each morph should track changes in the environment. Thus, facultative paedomorphosis may be maintained in part by variable, environmentally-specific fitness payoffs to each morph.     

Ernst Mayr as community architect: Launching the society for the study of evolution and the journalEvolution

Ernst Mayr's contributions to 20th century biology extend far beyond his defense of certain elements in evolutionary theory. At the center of mid-century efforts in American evolutionary studies to build large research communities, Mayr spearheaded campaigns to create a Society for the Study of Evolution and a dedicated journal,Evolution, in 1946. Begun to offset the prominence ofDrosophila biology and evolutionary genetics, these campaigns changed course repeatedly, as impediments appeared, tactics shifted, and compromises built a growing coalition of support. Preserved, however, were designs to balance the community and journal with careful equation of status and explicit partitioning of responsibilities within the working coalition. Choice terms such as cooperation and unity carried a strong political message. Mayr's editorship ofEvolution provides a superb example of these balancing efforts. The mid-century infrastructural activities described herein also represented aggressive attempts to leverage control across several layers of community. Leaders of these campaigns sought: (1) to promote evolutionary studies as a modernized research discipline and place it at the center of American biology, (2) to promote evolutionary studies within existing disciplines — e.g. systematics, genetics, and paleontology, (3) to foster certain research styles within evolutionary studies, and (4) to emphasize certain solutions to prominent research questions. Throughout, Mayr interjected his priorities, tactics and energy.     

The proximate/ultimate distinction in the multiple careers of Ernst Mayr

Ernst Mayr's distinction between “ultimate” and “proximate” causes is justly considered a major contribution to philosophy of biology. But how did Mayr come to this “philosophical” distinction, and what role did it play in his earlier “scientific” work? I address these issues by dividing Mayr's work into three careers or phases: 1) Mayr the naturalist/researcher, 2) Mayr the representative of and spokesman for evolutionary biology and systematics, and more recently 3) Mayr the historian and philosopher of biology. If we want to understand the role of the proximate/ultimate distinction in Mayr's more recent career as a philosopher and historian, then it helps to consider hisearlier use of the distinction, in the course of his research, and in his promotion of the professions of evolutionary biology and systematics. I believe that this approach would also shed light on some other important “philosophical” positions that Mayr has defended, including the distinction between “essentialism: and “population thinking.”     

The evolutionary expansion of the trypanosomatid flagellates

The trypanosomatids combine a relatively uniform morphology with ability to parasitise a very diverse range of hosts including animals, plants and other protists. Along with their sister family, the biflagellate bodonids, they are set apart from other eukaryotes by distinctive organisational features, such as the kinetoplast-mitochondrion and RNA editing, isolation of glycolysis enzymes in the glycosome, use of the flagellar pocket for molecular traffic into and out of the cell, a unique method of generating cortical microtubules, and bizarre nuclear organisation. These features testify to the antiquity and isolation of the kinetoplast-bearing flagellates (Kinetoplastida). Molecular sequencing techniques (especially small subunit ribosomal RNA gene sequencing) are now radically reshaping previous ideas on the phylogeny of these organisms. The idea that the monogenetic (MG) trypanosomatids gave rise to the digenetic (DG) genera is losing ground to a view that, after the bodonids, the African trypanosomes (DG) represent the most ancient lineage, followed by Trypanosoma cruzi (DG), then Blastocrithidia (MG), Herpetomonas (MG) and Phytomonas (DG), with Leptomonas (MG), Crithidia (MG), Leishmania (DG) and Endotrypanum (DG) forming the crown of the evolutionary tree. Vast genetic distances (12% divergence) separate T. brucei and T. cruzi, while the Leishmania species are separated by very short distances (less than 1% divergence). These phylogenetic conclusions are supported by studies on RNA editing and on the nature of the parasite surface. The trypanosomatids seem to be able to adapt with ease their energy metabolism to the availability of substrates and oxygen, and this may give them the ability to institute new life cycles if host behaviour patterns allow. Sexual processes, though present in at least some trypanosomatids, may have played only a minor part in generating diversity during trypanosomatid evolution. On the other hand, the development of altruistic behaviour on the part of some life cycle stages may be a hitherto unconsidered way of maximising fitness in this group. It is concluded that, owing to organisational constraints, the trypanosomatids can undergo substantial molecular variation while registering very little in the way of morphological change.     

Cytogenetical localization of Zygotic hybrid rescue (Zhr), a Drosophila melanogaster gene that rescues interspecific hybrids from embryonic lethality

Hybrid females from crosses between Drsophila melanogaster males and females of its sibling species, D. simulans, D. mauritiana, or D. sechellia die as embryos. This lethality is believed to be caused by incompatibility between the X chromosome of D. melanogaster and the maternal cytoplasm. Zygotic hybrid rescue (Zhr) prevents this embryonic lethality and has been cytogenetically mapped to a proximal region of the X chromosome of D. melanogaster, probably in the centromeric heterochromatin. We have carried out high resolution cytological mapping of Zhr using deficiencies and duplications of the X heterochromatin. Deletions of the Zhr ⁺ gene from the hybrid genome exhibit the Zhr phenotype. On the contrary, addition of the wild-type gene to the hybrid genome causes embryonic lethality, regardless of sex. The Zhr locus has been narrowed down to the region covered by Dp(1;f)1162 but not covered Dp(1;f)1205, a chromosome carrying a duplication of heterochromatin located slightly distal to the In(1)sc ⁸ heterochromatic breakpoint.     

Evolutionary epistemology as an overlapping,interlevel theory

I examine the branch of evolutionary epistemology which tries to account for the character of cognitive mechanisms in animals and humans by extending the biological theory of evolution to the neurophysiological substrates of cognition. Like Plotkin, I construe this branch as a struggling science, and attempt to characterize the sort of theory one might expect to find this truly interdisciplinary endeavor, an endeavor which encompasses not only evolutionary biology, cognitive psychology, and developmental neuroscience, but also and especially, the computational modeling of artificial life programming; I suggest that extending Schaffner's notion of interlevel theories to include both horizontal and vertical levels of abstraction best fits the theories currently being developed in cognitive science. Finally, I support this claim with examples drawn from computational modeling data using the genetic algorithm.     

Molecules and Morphology,Phylogenetics and Genetics

Various explanations can be offered for the incongruence between phylogenetic hypotheses resulting from morphological and molecular data sets. Of these, the possibility that incongruence may result from the mutation of major morphogenetic genes leading to dramatic morphological divergence unaccompanied by equivalent change of the phylogenetic marker molecule(s) used is discussed in detail. As evidence for this hypothesis, several examples for such incongruence are surveyed. It seems possible that in many cases the genetic basis of the morphological characters responsible for the incongruence found may be simple, and that the genes involved may be homologous to genes known from mutant systems. It is suggested that: 1. the systematic documentation of incongruence between molecular and morphological phylogenies may help to assess the frequency of evolutionary change through the mutation of major morphogenetic genes, and that 2. the identification of major morphological characters distinguishing closely related taxa with mutant phenotypes known from mutant systems eventually may allow an experimental approach to the problem of evolutionary change resulting from major genes. Natural taxa suspected to be the result of such processes could be changed morphologically through transformation with the relevant genes.     

Counterselection on sex chromosomes in the Mus musculus European hybrid zone

The extent to which alleles can disperse across a hybrid zone depends on the selection they are subjected to in the hybrid genetic background or, for those that are selectively neutral, on their ability to escape from the unfavourable environment by recombination. Three markers spanning a 45 cM segment in the center of the X chromosome were used to investigate the degree to which selection against X chromosome linked genes helps to maintain the barrier to gene flow in the hybrid zone between Mus musculus domesticus and M. m. musculus in Denmark. The introgression of all the sex chromosome specific markers was more limited than that of the autosomal enzymes (Idh1, Amy, Gpd1, Pgm1, Es1, Es2, Mpi, Np1, Es10, Sod1) and the mitochondrial DNA. The cline for DXPas2, which is in the center of the X chromosome, is extremely steep and shows that certain genes located in this region are strongly selected against in the hybrid background. The clines of the other two X-linked markers, Hprt and DXPas1, and of the Y chromosome are not as abrupt and all three have similar asymmetric introgression patterns. Although the musculus variants appear to behave in much the same way as those of the autosomal genes, the domesticus variants do not introgress. The results show that X-linked and to a lesser extent Y-linked genes are more strongly selected against in the hybrid genome than the mitochondrial genome or the different autosomal loci. This suggests that co-adapted gene systems involving the sex chromosomes may play an important role in the hybrid breakdown between the two subspecies.