Rachel Ben-Shlomo

The Evolutionary Significance of Genetic Diversity: Ecological, Demographic and Life History Correlates

The evolutionary significance of genetic diversity of proteins in nature remains controversial despite the numerous protein studies conducted electrophoretically during the last two decades. Ironically, the discovery of extensive protein polymorphisms in nature (reviewed by Lewontin, 1974; Powell, 1975; Selander, 1976; Nevo 1978, 1983b; Hamrick et al., 1979; Nelson and Hedgecock, 1980), did not resolve the disagreement between the die ho torn ou s explanatory models of selection (e.g., Ayala, 1977; Milkman, 1978; Clarke, 1979; Wills, 1981) versus neutrality (Kimura, 1968; Kimura and Chta, 1971; Nei, 1975; and modifications in Kimura, 1979atb). The more general problem of the relative importance of the evolutionary forces interacting in genetic population differentiation at the molecular levels of proteins and DNA, i.e., mutation, migration, natural selection and genetic drift, remains now as enigmatic as ever.

The genome sequence of the colonial chordate, Botryllus schlosseri

Botryllus schlosseri is a colonial urochordate that follows the chordate plan of development following sexual reproduction, but invokes a stem cell-mediated budding program during subsequent rounds of asexual reproduction. As urochordates are considered to be the closest living invertebrate relatives of vertebrates, they are ideal subjects for whole genome sequence analyses. Using a novel method for high-throughput sequencing of eukaryotic genomes, we sequenced and assembled 580 Mbp of the B. schlosseri genome. The genome assembly is comprised of nearly 14,000 intron-containing predicted genes, and 13,500 intron-less predicted genes, 40% of which could be confidently parceled into 13 (of 16 haploid) chromosomes. A comparison of homologous genes between B. schlosseri and other diverse taxonomic groups revealed genomic events underlying the evolution of vertebrates and lymphoid-mediated immunity. The B. schlosseri genome is a community resource for studying alternative modes of reproduction, natural transplantation reactions, and stem cell-mediated regeneration. DOI: http://dx.doi.org/10.7554/eLife.00569.001

Annual rhythms that underlie phenology: Biological time-keeping meets environmental change

Seasonal recurrence of biological processes (phenology) and its relationship to environmental change is recognized as being of key scientific and public concern, but its current study largely overlooks the extent to which phenology is based on biological time-keeping mechanisms. We highlight the relevance of physiological and neurobiological regulation for organisms’ responsiveness to environmental conditions. Focusing on avian and mammalian examples, we describe circannual rhythmicity of reproduction, migration and hibernation, and address responses of animals to photic and thermal conditions. Climate change and urbanization are used as urgent examples of anthropogenic influences that put biological timing systems under pressure. We furthermore propose that consideration of Homo sapiens as principally a ‘seasonal animal’ can inspire new perspectives for understanding medical and psychological problems.

Thermoregulatory and endocrine responses to light pulses in short-day acclimated social voles (Microtus socialis).

In mammals, nocturnal light pulses (NLP) have been demonstrated to affect physiology and behavior. However, the impact of NLP as a stressor has been less broadly examined. The purpose of this study was to examine the effect of NLP (three 15 min 450 lux light pulses) during each scotophase on both thermoregulation and endocrine stress responses under short‐day (SD; 8L:16D) acclimation. Voles were acclimated to either SD (SD voles) or SD+NLP (NLP voles). Resistance to cold was estimated by measurements of body temperature (Tb) during cold exposure (5°C). Daily rhythms of energy expenditure (calculated from oxygen consumption), urine production, and urinary adrenaline and serum cortisol levels were measured. Tb values of SD voles were generally unaffected by the cold stimulus, whereas in NLP voles, resistance to cold was markedly lowered. While SD‐ and NLP voles showed similar ultradian characteristics in energy expenditure with a period of 3.5 h, mean energy expenditure levels were lowest for voles exposed to NLP‐treatment. In SD voles, but not in NLP voles, urine production rates showed clear time variations and were consistently highest for SD voles, with significant differences during the scotophase. Both mean total urinary adrenaline and serum cortisol levels were significantly elevated in NLP‐treated voles compared with the control group. Taken together, the results suggest that NLP negatively affects winter acclimatization of thermoregulatory mechanisms of M. socialis, probably by mimicking summer acclimatization, and consequently the thermoregulatory mechanisms respond inappropriately to ambient conditions. One important finding of this study is that NLP may act as a stressor and correspondingly impose a major threat to the physiological homeostasis of M. socialis, such that over‐winter survival might be compromised.

Postglacial-period and Recent Invasions Shape the Population Genetics of Botryllid Ascidians along European Atlantic Coasts

The colonial urochordate Botryllus schlosseri is a sedentary species of Mediterranean origin that became cosmopolitan, probably because of postglacial-period dispersal and human-mediated invasions of colonies attached to ship hulls. Here we studied microsatellite allele diversity of Atlantic coast populations from an area ranging from European regions south of the last glacial front to regions that had been permanently ice-covered. Gene diversity levels varied dramatically among populations residing in areas subject to different glacial conditions. Five populations from the Iberian Peninsula, in an area south of the last glacial front, as well as two populations from presumed refugia in Brittany, expressed high gene diversity values (expected heterozygosity [He]: 0.76–0.80; average number of alleles per locus [A]: 7.25–8.75). Two populations inhabiting areas that experienced permafrost conditions (Helgoland Island, Germany, and Plymouth, England) had intermediate values (He: 0.40–0.42; A: 3.0–4.0), whereas the Auchenmalg, Scotland, population, from an area previously covered by ice, showed a remarkably low value (He: 0.17; A: 1.75). Therefore, most European populations of B. schlosseri mirrored the movement of the ice front in the last ice age. A second population from the area that was covered by permanent ice (Lossiemouth, Scotland), however, had a high He of 0.61 and an intermediate A of 3.67. Results were compared with recent invasions (populations less than 200 years old) in the United States and New Zealand that had a higher degree of genetic variation than the European native populations established thousands of years ago. Given the overall dearth of studies on this subject, we suggest that in contemporary established Botryllus populations, gene diversity is affected by ecological factors, some of which can be traced directly to the last ice age. Other parameters of gene diversity are influenced by selection pressure, which might be more intense in northern regions.

Activity pattern and rhythm in the subterranean mole rat superspecies Spalax ehrenbergi.

Good candidates for naturally occurring variability in circadian rhythms may be subterranean herbivores, since they are not normally subjected to entraining light stimulation. To test this possibility, we selected the blind mole ratSpalax ehrenbergi superspecies in Israel and tested it in short- and long-term experiments. Short-term experiments showed that the animals exhibited three patterns of activity: a regular circadian rhythm (26.6%), an altered circadian rhythm (shorter or longer than normal, 53.1%), and an arrhythmic pattern (20.3%). A long-term experiment showed that the arrhythmic pattern indeed reflected a genuine arrhythmic genotype. The mole rats were found to be active less than 25% of the day and exhibited a multiphasic mode of activity, both diurnally and nocturnally. The number of activity periods and the level of activity were negatively correlated: Animals that exhibited a high level of activity per unit of time showed low numbers of activity periods, while animals that exhibited a lower level of activity showed higher numbers.

Isozyme polymorphism as monitoring of marine environments: The interactive effect of cadmium and mercury pollution on the shrimp, Palaemon elegans

Abstract The separate and interactive pollution effects of cadmium and mercury on the electrophoretically detected allozyme frequencies of the phosphoglucomutase isozyme were tested in Palaemon elegans . Our results indicate differential survivorship of allozyme genotypes specific for each metal and for their interaction. These results reflect the adaptive nature of the allozyme genotypes tested in this shrimp and seem to be inconsistent with the neutral theory of allozyme polymorphisms. Furthermore, these findings suggest that the interactive contamination of cadmium and mercury acts as a specific contaminant. Therefore, differential tolerance of allozyme genotypes to metals can be used as a detector of heavy metal contamination in the sea not only for individual pollutants, but also for many pollutants and their interactive effects.

Pattern of settlement and natural chimerism in the colonial urochordate Botryllus schlosseri

Colonies of the cosmopolitan urochordate Botryllus schlosseri that share one or both alleles at a single allorecognition locus (Fu/HC) and come into tissue contacts, may fuse and form a mixed entity, a chimera. Botryllus populations worldwide exhibit unprecedented extensive polymorphism at this locus, a result that restricts fusions to kin encounters. This study aims to compare spatiotemporal configurations in source and introduced B. schlosseri populations, residing on natural and man-made substrata, respectively. By using four microsatellite loci, we tested genetic consanguinity of colonies settled naturally along spatial vectors on both, natural (native populations) and man-made (introduced) substrates. Four populations were studied. Results revealed that B. schlosseri colonies, on both substrate types, assemble in groups of relatives that share similar microsatellite profiles. We suggest that this pattern of settlement promotes the formation of chimeras, which evoke conflicting interactions: cooperation between different somatic cell lines that constitute the colonial soma and competition between germ cells that inhabit the chimera gonads. Under natural conditions, the chimera may allow genetic flexibility that depends on joint genomic fitness of its partners. This is probably one of the life history characteristics that led to the worldwide distribution success of this species.

Allozyme differentiation and systematics of the endemic subterranean mole rats of South Africa

Abstract We have analyzed allozyme differentiation, encoded by 20 putative gene loci, of three genera and four currently accepted species of Southern African mole rats, Bathyergidae ( Cryptomys hottentotus hottentotus, C. hottentotus natalensis, C. hottentotus damarensis, Georychus capensis, Bathyergus suillus and B. janetta ). Two major results are indicated: (a) genetically the family reflects distinct homozygosity ( H =0.019, range 0.00–0.05) as is typical for underground fossorial mammals and (b) remarkable large genetic distances exist between the six taxa analyzed ( D = 1.865, range 0.547–2.917). These results suggest that homoselection operates in the underground ecotope in accordance with the niche width variation hypothesis, and that bathyergid systematics needs urgent revision, particularly the genus Cryptomys . The three subspecies of Cryptomys appear to be at least three different species, possibly even falling into two genera. The adaptive radiation of bathyergids started at least in Miocene times, but probably began earlier in the Oligocene as indicated by the fossil record.

Haptoglobin DNA polymorphism in subterranean mole rats of the Spalax ehrenbergi superspecies in Israel

We analyzed the genetic diversity and environmental correlates of the haptoglobin (Hp) gene by RFLP analysis of 121 subterranean mole rats, comprising 13 populations belonging to the 4 chromosomal species (2n = 52, 54, 58 and 60) of the actively speciating Spalax ehrenbergi superspecies in Israel. The following results were indicated: (i) Out of the six restriction enzymes we tested, we found polymorphism only in TaqI, with three allelic fragments, and described their geographic patterning, (ii) Genetic diversity of Hp was highest in the centre of the range, (iii) The two major alleles showed high significant and opposite correlation with humidity, (iv) Genetic differentiation analysis indicated that 40 per cent of the Hp variation was within, and 60 per cent between populations. We conclude that climatic selection plays an important role in haptoglobin genetic differentiation.