The carbonate factory continuum, facies mosaics and microfacies: an appraisal of some of the key concepts underpinning carbonate sedimentology

Our understanding of where and how carbonate sediments are produced and accumulate has changed considerably in recent years and a more complex framework is emerging. The earlier concept invoking a limited range of productivity-depth models has now evolved into an appreciation that there is a continuum of different types of productive sites over wide depth ranges, influenced by complex factors and not simply water depth or temperature. Studies of the nature of lithofacies ordering in the stratigraphic record, and most recent studies of the spatial distribution of Holocene environments, raise the issue that at the lithofacies scale the sedimentary record represents, in part, the product of complex and mobile facies mosaics. Many of these mosaic elements are not depth dependent and can change through time as a consequence of subtle environmental changes. As the rates of change typically exceed rates of accommodation space creation, individual sites are likely to have sediments of different environments superimposed and mixed (palimpsest). Recent studies showing the extent that dissolution is capable of skewing sediment compositions suggest that many ancient microfacies are unrepresentative of their original sediments, and there is a need for a more critical approach to interpreting microfacies in terms of identifying habitats and especially water depth. The carbonate factory is spatially and temporally highly variable and is not simply a uniform production line. This fact, coupled with the likely importance of selective early dissolution, may in part explain why accumulation rates estimated from ancient strata are lower than the production rates measured over short time periods.     

Marine carbonate facies in response to climate and nutrient level: The upper carboniferous and permian of central spitsbergen (Svalbard)

Summary Carbonate-dominated successions of the Gipsdalen and Tempelfjorden Groups from Svalbard record a significant shift from Photozoan to Heterozoan particle associations in neritic settings during the late Palaeozoic. During the Bashkirian, benthic particle associations which included photoautotrophs such as phylloid algae (Chloroforam Association) characterised shallow subtidal environments. Most depositional settings which endured siliciclastic terrestrial input exhibited poorly diversified associations dominated by brachiopods, bryozoans and siliceous sponges (Bryonoderm Association). During the Moscovian to Asselian, highly diversified associations typified by various calcareous green algae,Palaeoaplysina, Tubiphytes, fusulinids, smaller and encrusting foraminifers (Chloroforam Association) prevailed in carbonate sediments from supratidal to shallow subtidal environments. During the Sakmarian and Early Artinskian, oolitic carbonate sands (Chloroforam Association) typified intertidal flats, whereas shallow subtidal environments were occupied by moderately diversified associations with fusulinids, smaller foraminifers, echinoderms and bryozoans (Bryonoderm-extended Association) and poorly diversified associations with echinoderms, brachiopods and bryozoans (Bryonoderm Association). During the Late Artinskian to Kazanian, poorly diversified associations characterised by brachiopods, echinoderms and bryozoans (Bryonoderm Association), and sponge-dominated associations (Hyalosponge Association) reigned within siliceous carbonates of intertidal and shallow subtidal environments. This trend is interpreted as a result of climatic cooling and fluctuations of prevailing levels of trophic resources within shallow-water settings during the studied time period. While raised nutrient levels were restricted to near-shore settings during the Bashkirian, steady mesotrophic conditions arose from the Sakmarian onward and increased to late Permian times.     

Foraminifers of the Viséan–Serpukhovian boundary interval in Western Palaeotethys: a review

A detailed revision of foraminiferal zonal schemes in sections throughout Europe and North Africa for the Viséan–Serpukhovian boundary interval suggests that several foraminiferal taxa might have the potential to form reliable markers throughout the Palaeotethys. This would support the currently investigated boundary definition based on the First Appearance Datum of the conodont Lochriea ziegleri. However, correlation of these foraminiferal markers in the Western Palaeotethys region has encountered several problems, partly arising from taxonomic issues, but mainly because of apparent discrepancies between the First Occurrence Data (FOD). Analysis of the available foraminiferal data has revealed that some taxa show marked delays in their FODs, due to the timing of westward dispersal within the Palaeotethys, emanating from a probable source in eastern Russia. As a result of this investigation, two dispersal routes have been identified, a northern branch and a southern branch. In general, the displacements within the southern branch occurred more rapidly than in the northern branch. In addition to different dispersal routes, separation of the main foraminiferal markers in stratigraphical sections from different regions can result from isolation of shallow‐water facies of the inner platform from those of relatively deeper‐water settings in the outer platform, the latter showing more consistent foraminiferal FODs. The differences in palaeobathymetry and associated energy levels have enabled two foraminiferal zonal schemes to be distinguished for the Viséan–Serpukhovian boundary interval in the Western Palaeotethys, one for the inner platform and a second one for the outer platform.     

Progressive drowning of carbonate platform in the Moravo-Silesian Basin (Czech Republic) before the Frasnian/Famennian event: facies,compositional variations and gamma-ray spectrometry

The Moravo-Silesian Basin (MSB; eastern Czech Republic and southern Poland) hosted an extensive shallow-water carbonate platform in the Middle Devonian to Frasnian interval. The platform drowned in a stepwise fashion from the Palmatolepis hassi to the Pa. linguiformis zone. Three types of drowning successions were revealed from conodont biostratigraphy, facies, microfacies and gamma-ray spectrometry data: (A) drowning to periplatform turbidite setting; (B) drowning to (hemi)pelagic seamount setting and (C) drowning associated with the stratigraphical gap. In the lower Pa. hassi zone, rapid subsidence caused the platform to drown locally along the N–S to NW–SE trending faults (type A drowning). In the upper Pa. rhenana to the Pa. linguiformis zone, the drowning accelerated in the western part of the MSB due to locally higher subsidence rates combined with the Late Frasnian biotic crisis (type B). In the southern part of the basin, the platform emerged shortly before the Frasnian/Famennian (F/F) boundary and drowned in the Early to Late Famennian (type C). The primary cause of drowning was differential subsidence at the Laurussian passive margin. Eustatic sea-level fluctuations, if any, contributed only to a minor extent to the Late Frasnian drowning, but were effective in type C drowning during the Famennian. The drowning boundaries are associated with increased contents of K and Th, reflecting the deceleration of carbonate production. Uranium contents display isolated peaks that roughly correlate with the drowning boundaries or the stratigraphic gaps associated with the F/F boundary. The uranium contents are considered to reflect local depositional conditions and are not suitable for stratigraphic correlation. On the other hand, from the K and Th contents, we can infer Late Frasnian sea-level fluctuations with duration on the order of 1 Myr. These cyclic variations in K and Th contents proved to be useful in platform-to-basin stratigraphic correlation.     

Allozyme differentiation and relationships within Hypostomus Lacépède, 1803 (Osteichthyes: Loricariidae) from the upper Paraguay River basin, Brazil

Nine hypostomine populations from Manso Reservoir (Rio Manso and Rio Cuiabá, upper Paraguay River basin, Brazil) were surveyed using allozyme electrophoresis. Three, sampled at the same sites, were not identified to the species level and were designated as Hypostomus sp. 1, Hypostomus sp. 2 and Hypostomus sp. 3. The survey resulted in 25 loci from 14 scored enzyme systems. Allozyme data allowed diagnostic marker recording for all species analyzed, including the undescribed taxa. Mean expected heterozygosity values were quite variable, ranging from 0.029 to 0.079. Hypotheses to explain this variation are discussed. The allelic frequencies and Nei's genetic identity values showed that Hypostomus sp. 1, Hypostomus sp. 2 and Hypostomus sp. 3 should be considered true biological species, and distinct from the other five Hypostomus species and Pterygoplichthys anisitsi.     

Devonohexapodus bocksbergensis is a synonym of Wingertshellicus backesi (Euarthropoda) – no evidence for marine hexapods living in the Devonian Hunsrück Sea

The Devonian Hunsrück Slate fossil Devonohexapodus bocksbergensis Haas, Waloszek & Hartenberger, 2003 has been interpreted as a stem-lineage representative of the Hexapoda, implying their marine origin and independent terrestrialisation within the ‘Atelocerata’. Devonohexapodus bocksbergensis was based on a single specimen embedded in a lateral position. Reinvestigation of that holotype and of all known specimens of the Hunsrück Slate arthropod Wingertshellicus backesi Briggs & Bartels, 2001 demonstrates that all this material represents a single species. The latter is redescribed, its taxonomic diagnosis is emended, and the name Devonohexapodus bocksbergensis is treated as a junior synonym of Wingertshellicus backesi. The phylogenetic position of W. backesi neither is that of a stem-lineage representative of Hexapoda, nor does it fall within the crown group Mandibulata. The Hunsrück Slate provides no evidence of an independent terrestrialisation within the ‘Atelocerata’ or of a marine origin of the Hexapoda.     

The arachiform vacuolar body: an overlooked shared character in the ascospores of a large monophyletic group within Parmeliaceae (Xanthoparmelia clade, Lecanorales)

Sections of apothecia were used to study the internal morphology of ascospores in the largest monophyletic clade within Parmeliaceae composed of Xanthoparmelia and related genera. The results were compared with fertile representative species of most other parmelioid clades. All the Xanthoparmelia species had spores with a single smooth vacuole, which was peanut-shaped, with different degrees of constriction in the equatorial plane. This differs from the ellipsoid vacuole of other parmelioids. In the Xanthoparmelia clade, sexual reproduction seems much more common than in other parmelioids. Thus, we suggest that the presence of this unique spore morphology might contribute to the evolutionary success of this monophyletic group. Further, the discovery of this useful ascospore character demonstrates that detailed ascospore morphological studies significantly enhance molecular phylogenetic analyses. Ascospore features may be more taxonomically significant in Parmeliaceae than hitherto considered.     

Studies on the continuous production of (R)-(−)-phenylacetylcarbinol in an enzyme-membrane reactor

The optimization of a continuous enzymatic reaction yielding (R)-(−)-phenylacetylcarbinol ((R)-PAC), a key intermediate of the (1R,2S)-(−)-ephedrine synthesis, is presented. We compare the suitability of different mutants of the pyruvate decarboxylase (PDC) from Zymomonas mobilis with respect to their application in biotransformation using pyruvate or acetaldehyde and benzaldehyde as substrates, respectively. Starting from 90 mM pyruvate and 30 mM benzaldehyde, (R)-PAC was obtained with a space time yield of 27.4 g/(L·day) using purified PDCW392I in an enzyme-membrane reactor. Due to the high stability of the mutant enzymes PDCW392I and PDCW392M towards acetaldehyde, a continuous procedure using acetaldehyde instead of pyruvate was developed. The kinetic results of the enzymatic synthesis starting from acetaldehyde and benzaldehyde demonstrate that the carboligation to (R)-PAC is most efficiently performed using a continuous reaction system and feeding both aldehydes in equimolar concentration. Starting from an inlet concentration of 50 mM of both aldehydes, (R)-PAC was obtained with a space-time yield of 81 g/(L·day) using the mutant enzyme PDCW392M. The new reaction strategy allows the enzymatic synthesis of (R)-PAC from cheap substrates free of unwanted by-products with potent mutants of PDC from Z. mobilis in an aqueous reaction system.     

The importance of microhabitat in thermoregulation and thermal conductance in two namib rodents—a crevice dweller, Aethomys namaquensis, and a burrow dweller, Gerbillurus paeba

1. 1.|Thermoregulatory measurements of two Nambi rodents; Gerbillurus paeba, a burrow dweller, and Aethomys namaquensis, a crevice dweller were compared. Both were similar to other small arid-adapted rodents in that basal metabolic rates were reduced, thermoneutral zones narrow and evaporative water losses low. Rates of conductance and thermal lability, however, at ambient temperatures (T_a) below thermoneutral zone, were significantly different (P 0.01).

2. 2.|The rock rat A. namaquensis, living in a microclimate characterized by a large diel range and low humidities, compensates for a reduced basal metabolic rate by having a low rate of conductance. In this way it maintains precise thermoregulatory control. G. paeba, on the other hand, living in a thermally-stable milieu, does not control body temperature precisely. This animal instead utilizes a high rate of conductance to remove metabolic heat produced within the body. This would be advantageous to an animal living in a plugged burrow where the high humidities encountered impede the rate of evaporative cooling.

3. 3.|The energetic responses of both species, above the thermoneutral zone, appear to reflect very closely the environmental conditions which occur in the microhabitat that they rest in during the day. G. paeba shows less tolerance to temperature fluctuations than A. namaquensis, but shows more marked increases in short-term cooling mechanisms at high T_as.

4. 4.|Despite the increased use of evaporative cooling through salivation and panting in addition to pulmocutaneous evaporation, exposure to T_as above 38°C is rapidly lethal to G. paeba.

The presence of Tawera gayi (Hupé in Gay, 1854) (Veneridae, Bivalvia) in southern South America: Did Tawera achieve a Late Cenozoic circumpolar traverse?

Cenozoic Tawera Marwick, 1927 from the Southern Hemisphere exhibits a pattern of disjunt distribution around the southern oceans. A single species, Tawera gayi (Hupé in [Gay, C. (1854). Historia Física y Política de Chile, Zoología 8. Paris.]) is confined to southern South America. Taking into account the occurrence of Tawera in the fossil record, taxonomy based on shell morphology, and available information on extant species of Tawera, it is plausible that the genus appeared first in southern Australia during the Early Miocene, and then expanded and radiated to New Zealand. It also appears that Tawera first crossed from Australasia to South America during the Early Pleistocene. This picture can be better explained if Tawera was able to achieve circumglobal range by means of the Antarctic Circumpolar Current. Thus, different potential factors of dispersal (i.e., larval dispersal, drifting, kelp rafting and Pleistocene cooling) are considered and discussed.

Shell morphology and overall appearance of Tawera gayi is very similar to Tawera philomela (Smith, 1885) from South Africa and Tawera mawsoni (Hedley, 1916) from Macquarie Island, suggesting these taxa have a close relationship. One postulated explanation, which should be confirmed by means of a phylogenetic study, is a subsequent migration of Tawera from South America arriving first to the Southern African Region (via the West Wind Drift Islands Province), and then probably coming back again to Australasia. It could have been mediated via the same current during the Late Pleistocene and much later during the Holocene.     

Rice chloroplast RNA polymerase genes: The absence of an intron in rpoC1 and the presence of an extra sequence in rpoC2

Summary The chloroplast genome contains sequences homologous to the Escherichia coli rpoA, rpoB and rpoC genes. The Choroplast rpoC gene is divided into rpoC1 and rpoC2, of which rpoC1 contains an intron. Comparison of the rice rpo genes with those from tobacco, spinach and liverwort revealed unique features of the rice genes; the lack of an intron in rpoC1 and the presence of an extra sequence of 381 by in rpoC2. The intron in rpoC1 is thus optional, and possible intron boundary sites in split rpoC1 genes can be estimated by comparison with rice rpoC1. The extra sequence is located in the middle of rpoC2 and has repeated structures. The amino acid sequence deduced from this sequence is extremely hydrophilic and anionic. The origin and function of this sequence are discussed.     

The organization of the chemosensory system in Drosophila melanogaster: a rewiew

This review surveys the organization of the olfactory and gustatory systems in the imago and in the larva of Drosophila melanogaster, both at the sensory and the central level. Olfactory epithelia of the adult are located primarily on the third antennal segment (funiculus) and on the maxillary palps. About 200 basiconic (BS), 150 trichoid (TS) and 60 coeloconic sensilla (CS) cover the surface of the funiculus, and an additional 60 BS are located on the maxillary palps. Males possess about 30% more TS but 20% fewer BS than females. All these sensilla are multineuronal; they may be purely olfactory or multimodal with an olfactory component. Antennal and maxillary afferents converge onto approximately 35 glomeruli within the antennal lobe. These projections obey precise rules: individual fibers are glomerulus-specific, and different types of sensilla are associated with particular subsets of glomeruli. Possible functions of antennal glomeruli are discussed. In contrast to olfactory sensilla, gustatory sensilla of the imago are located at many sites, including the labellum, the pharynx, the legs, the wing margin and the female genitalia. Each of these sensory sites has its own central target. Taste sensilla are usually composed of one mechano-and three chemosensory neurons. Individual chemosensory neurons within a sensillum respond to distinct subsets of molecules and project into different central target regions. The chemosensory system of the larva is much simpler and consists essentially of three major sensillar complexes on the cephalic lobe, the dorsal, terminal and ventral organs, and a series of pharyngeal sensilla.     

Horizontal transmission of a microsporidium from the convergent lady beetle, Hippodamia convergens Guérin-Méneville (Coleoptera: Coccinellidae), to three coccinellid species of Nova Scotia

Convergent lady beetles, Hippodamia convergens Guérin-Méneville, are a popular choice for aphid control in North America. An unidentified microsporidium was found in H. convergens adults that were purchased from a commercial insectary in 2004. This study examined egg cannibalism and egg predation as a means of horizontal transmission of the unidentified microsporidium among H. convergens larvae and three coccinellid species found in Nova Scotia: Coccinella septempunctata (seven-spotted lady beetle), C. trifasciata perplexa (three-banded lady beetle), and Harmonia axyridis (multicolored Asian lady beetle). The microsporidium was transmitted with 100% efficiency when first instars fed on microsporidia-infected eggs. Mean spore count data from smear preparations of infected beetles suggest that the infection was as heavy in C. trifasciata perplexa (a native coccinellid) (11.2 ± 0.96 spores/100 μm²) as it was in H. convergens (the natural host) (12.8 ± 1.16) but lighter in the introduced species C. septempunctata (7.5 ± 0.65) and H. axyridis (0.8 ± 0.11). For all of the beetle species examined, larval development was significantly longer for microsporidia-infected individuals than for their uninfected cohorts. The microsporidium had no effect on larval mortality. Based on the results of this study, field-collected H. convergens should be examined for microsporidia and uninfected individuals should be used to rear individuals for release in biological control programs. However, this is unlikely to happen because H. convergens are relatively easy and inexpensive to collect from their overwintering sites for redistribution.     

The impact of temperature on the inactivation of enteric viruses in food and water: a review

Temperature is considered as the major factor determining virus inactivation in the environment. Food industries, therefore, widely apply temperature as virus inactivating parameter. This review encompasses an overview of viral inactivation and virus genome degradation data from published literature as well as a statistical analysis and the development of empirical formulae to predict virus inactivation. A total of 658 data (time to obtain a first log(10) reduction) were collected from 76 published studies with 563 data on virus infectivity and 95 data on genome degradation. Linear model fitting was applied to analyse the effects of temperature, virus species, detection method (cell culture or molecular methods), matrix (simple or complex) and temperature category (<50 and ≥50°C). As expected, virus inactivation was found to be faster at temperatures ≥50°C than at temperatures <50°C, but there was also a significant temperature-matrix effect. Virus inactivation appeared to occur faster in complex than in simple matrices. In general, bacteriophages PRD1 and PhiX174 appeared to be highly persistent whatever the matrix or the temperature, which makes them useful indicators for virus inactivation studies. The virus genome was shown to be more resistant than infectious virus. Simple empirical formulas were developed that can be used to predict virus inactivation and genome degradation for untested temperatures, time points or even virus strains.     

Immunocytochemical study of tubulin in the 9+‘1’ sperm axoneme of a monogenean (Platyhelminthes), Pseudodactylogyrus sp.

The spermatozoon of the monopisthocotylean monogenean Pseudodactylogyrus sp. (a gill parasite of eels) has a single axoneme showing a 9+‘1’ pattern, a nucleus and a mitochondrion, but has no cortical microtubules. This species thus provides a very simple model for the study of tubulin in the 9+‘1’ axonemes of the Platyhelminthes, in contrast with digenean sperm which have a more complex spermatozoon with two such axonemes and cortical microtubules. Indirect immunofluorescence labelling of tubulin shows that the elongating spermatids, initially lying in all directions in the early stages, are arranged as parallel elements in further stages. The number of spermatids in an isogenic group could also be precisely counted and equals 32. Nuclear labelling with fluorescent dyes shows that the nuclei, first located in the common mass of the spermatids, later elongate and migrate into the growing spermatids, and that the nucleus is located in the central part of the mature spermatozoon, with the two extremities devoid of nucleus. Labelling with antibodies directed against acetylated, tyrosinated, and polyglutamylated tubulin gave positive results, thus indicating that these post-translational modifications of tubulin are present in the axoneme of spermatids and spermatozoa of monopisthocotylean monogeneans.     

Transformation of mouse mammary epithelial cells with the Ha-ras but not with the neu oncogene results in a gene dosage-dependent increase in transforming growth factor-α production

An enhanced expression of transforming growth factor-α (TGFα) was demonstrated in two clones of NOG-8 mouse mammary epithelial cells, NOG-8 SR1 and NOG-8 SR2, that have been transformed by a v-Ha-ras oncogene. The amount of TGFα production in NOG-8 SR1 and NOG-8 SR2 cells was dependent on the level of p21^ras expression in these clones, which directly correlated with their cloning efficiency in soft agar. There was also a decrease in the number of epidermal growth factor (EGF) receptors on the NOG-8 SR1 and NOG-8 SR2 cells that is proportional to the amount of TGFα secreted. These effects were specific for ras because neu-transformed NOG-8 cells grew in soft agar at a comparable level to NOG-8 SR2 cells yet did not show any increase in TGFα production or change in EGF receptor expression.     

The free reducing oligosaccharides of angico branco (Anadenanthera colubrina) gum exudate: an aid for structural assignments in the heteropolysaccharide

A novel method is described for the determination of sequential side-chain structures in the complex, high-arabinose polysaccharide of the gum exudate of angico branco (Anadenanthera colubrina), using as basis the structurally similar reducing oligosaccharides present in small quantities. Of the ten detected, eight were characterized as disaccharides (2, 3, and 9), linear trisaccharides (1 and 4), branched pentasaccharides (5 and 6), and a doubly branched heptasaccharide (8). The oligosaccharides are substituents of the polysaccharide, which has a (1→3)-linked β- -galactopyranosyl main chain, and with two exceptions they had 6-O-substituted galactopyranosyl reducing ends, probably corresponding to its main-chain units. Characterization was effected through their 1D and 2D NMR correlation spectra, which were better resolved and more readily interpretable than those of the polysaccharide. These spectral data were supported by monosaccharide composition and rotation values. Controlled Smith degradations and methylation analyses were carried out when it was necessary. These data were confirmed by field-desorption MS.     

Spacing patterns of an Acacia tree in the Kalahari over a 61-year period: How clumped becomes regular and vice versa

Nearest tree neighbour distances and the tree spatial formation on a large scale over time and space replicates were examined. The study was conducted in a natural savanna ecosystem in the Southern Kalahari, South Africa. Nearest tree neighbour and point pattern analysis methods were used to investigate changes in the spatial pattern of trees in two plots. Trees larger than 2 m canopy diameter were mapped. We used aerial photographs of the study area from 1940, 1964, 1984, 1993, and a satellite image from 2001 to follow two plots over time. Field work was carried out too for classification accuracy. We were able to identify and individually follow over 2400 individual trees from 1940 until 2001. Nearest neighbour analysis results indicate that dead trees were on average closer to their nearest neighbouring trees than living trees were to their neighbours. Most dead trees were on average 6 m from their nearest neighbours, while most living trees were about 20 m apart. Point pattern analysis results show a cyclical transition from clumped to random and sequentially to regular tree spacing. These transitions were not correlated across two plots. Generally, decreases in small-scale clumping coincided with periods of high mortality. Our findings show that regular, clumped, and random tree pattern can occur, pending on time, location, and scale within the location.     

DNA sequence analysis of spontaneous and γ-radiation (anoxic)-induced lacI mutations in Escherichia coliumuC122::Tn5: differential requirement for umuC at G · C vs. A · T sites and for the production of transversions vs. transitions

Escherihica coliumC122::Tn5 cells were γ-radiated (¹³⁷Cs, 750 Gy, under N₂), and lac-constitutive mutants were produced at 36% of the wild-type level (the umC strain was not deficient in spontaneous mutagenesis, and the mutational spectrum determined by sequencing 263 spontaneous lacI^d mutations was very similar to that for the wild-type strain). The specific nature of the umC strain's partial radiation was determined by sequencing 325 radiation-induced lacI^d mutations. The yields of radiation-induced mutation classes in the umC strain (as a percentage of the wild-type yield) were: 80% for A · T → G · C transitions, 70% for multi-base additions, 60% for single-base deletions, 53% for A · T → C · G transversions, 36% for G · C → A · T transitions, 25% for multi-base deletions, 21% for A · T → T · A transversions, 11% for G · C → C · G transversions, 9% for G · C → T · A transversions and 0% for multiple mutations. Based on these deficiencies and other factors, it is concluded that the umC strain is near-normal for A · T → G · C transitions, single-base deletions and possibly A · T → C · G transversions; is generally deficient for mutagenesis at G · C sites fro transversions, and is grossly deficient in multiple mutations. Damage at G · C sites seems more difficult for translesion DNA synthesis to bypass than damage at A · T sites, and especially when trying to produced a transversion. The yield of G · C → A · T transitions in the umC strain *36% of the wild-type level) argues that a basic sites are involved in no more than 64% of γ-radiation-induced base substitutions in the wild-type strain. Altogether, these data suggest that the UmuC and UmuD′ proteins facilitate, rather than being absolutely required for, translesion DNA synthesis; with the degree of facilitation being dependent both on the nature of the noncoding DNA damage, i.e., at G · C vs A · T sites, and on the nature of the misincorporated base, i.e., whether it induces transversions or transitions.