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Geographic ranges of species and their determinants are of great interest in the field of biogeography and are often studied in terms of the species" ecological niches. In this context, the range of a species is defined by the accessibility of an area, abiotic factors and biotic interactions, which affect a species" distributions with different intensities across spatial scales. Parapatry describes a distributional pattern in which the ranges of two species meet along sharp range limits with narrow contact zones. Such parapatric range limits are determined by changing abiotic conditions along sharp environmental gradients or can result from interspecific resource competition. However, it has been shown that often the interplay of abiotic conditions and species interactions determine parapatry. The geographic ranges of the land salamanders, Salamandra salamandra and S. atra, narrowly overlap in the European Alps with only few syntopic localities and to date, the cause of parapatry is unknown. The goal of this thesis was thus to identify the importance of abiotic and biotic factors for their parapatric range limits at different spatial scales. On a broad spatial scale, the role of climate for the parapatric range limits of the species was investigated within three contact zones in Switzerland. Climatic conditions at species" records were analysed and species distribution modelling techniques were used to explore the species" climatic niches and to quantify the interspecific niche overlap. Furthermore, it was tested whether the parapatric range limit coincides with a strong climatic gradient. The results revealed distinct niches for the species as well as the presence of strong climatic gradients which could explain the parapatric range limits of the species. Yet, there was a moderate interspecific niche overlap in all contact zones indicating that the species may co-occur and interact with each other in areas where they both find adequate conditions. Comparison among contact zones revealed geographic variation in the species" niches as well as in the climatic conditions at their records suggesting that the species can occur in a much wider range of conditions than they actually do. These findings imply that climate represents a main factor for the species" parapatric range limits. Yet, interspecific niche overlap and the geographic variation provide indirect evidence that interspecific interaction may also affect their spatial distribution. To test whether competition restricts the species" ranges on the habitat scale and to understand local syntopic co-occurrence of the salamanders within their contact zones, site-occupancy modelling was used. This approach allowed to find the habitat predictors that best explain the species" local distribution. While the slope of the site positively affected the occupancy probability of S. salamandra, no tested predictor explained that of S. atra. Also, there was no effect of the occurrence of one species on the occupancy probability of the other providing no evidence for competition. Should competition occur, it does not lead to spatial segregation of the species on this scale. Because biotic interactions most significantly affect the ranges of species on small spatial scales, the microhabitat conditions at locations of the species within syntopic contact zones were compared and a null model analysis was applied to determine their niche overlap. Resource selection probability function models were used to assess those attributes that affect the species" habitat selections. The results revealed species-specific microhabitat preferences related to leaf litter cover, tree number and that the species were active at different temperatures as well as times of the day. The high degree of diurnal activity of S. atra may be due to its preference of forest floor microhabitats that long remain suitable during daytime. Besides, there was a great niche overlap for shelters indicating that the species may compete for this resource. Differential habitat selection and the use of the available shelters at different times of the day may minimize species interactions and allow their local co-occurrence within contact zones. To identify whether the potential infection with the pathogenic chytrid fungus could serve as an alternative biotic explanation for the range margins of S. atra, several populations throughout its range were screened for infection. Since the occurrence of this pathogen was detected mostly at lower altitudes of the Alps, it may confine the range of S. atra to higher elevations. Because chytrid was not detected in any of the samples, the pathogen unlikely plays a role in determining its range limits. Overall, these findings underline the complexity of mechanisms that determine the range margins of parapatric species and provide an important basis for subsequent studies regarding the determinants of the parapatric distribution of the two salamander species.
In this thesis, in order to shed light on the biological function of the membrane-bound Glucocorticoid Receptor (mGR), proteomic changes induced by 15 min in vivo acute stress and by short in vitro activation of the mGR were analyzed in T-lymphocytes. The numerous overlaps between the two datasets suggest that the mGR mediates physiologically relevant actions and participates in the early stress response, triggering rapid early priming events that pave the way for the slower genomic GC activities. In addition, a new commercially available method with suitable sensitivity to detect the human mGR is reported and the transcriptional origin of this protein investigated. Our results indicates that specific GR-transcripts, containing exon 1C and 1D, are associated with the expression of this membrane isoform.
The availability of data on the feeding habits of species of conservation value may be of great importance to develop analyses for both scientific and management purposes. Stomach flushing is a harmless technique that allowed us to collect extensive data on the feeding habits of six Hydromantes species. Here, we present two datasets originating from a three-year study performed in multiple seasons (spring and autumn) on 19 different populations of cave salamanders. The first dataset contains data of the stomach content of 1,250 salamanders, where 6,010 items were recognized; the second one reports the size of the intact prey items found in the stomachs. These datasets integrate considerably data already available on the diet of the European plethodontid salamanders, being also of potential use for large scale meta-analyses on amphibian diet.
Species can show strong variation of local abundance across their ranges. Recent analyses suggested that variation in abundance can be related to environmental suitability, as the highest abundances are often observed in populations living in the most suitable areas. However, there is limited information on the mechanisms through which variation in environmental suitability determines abundance. We analysed populations of the microendemic salamander Hydromantes flavus, and tested several hypotheses on potential relationships linking environmental suitability to population parameters. For multiple populations across the whole species range, we assessed suitability using species distribution models, and measured density, activity level, food intake and body condition index. In high-suitability sites, the density of salamanders was up to 30-times higher than in the least suitable ones. Variation in activity levels and population performance can explain such variation of abundance. In high-suitability sites, salamanders were active close to the surface, and showed a low frequency of empty stomachs. Furthermore, when taking into account seasonal variation, body condition was better in the most suitable sites. Our results show that the strong relationship between environmental suitability and population abundance can be mediated by the variation of parameters strongly linked to individual performance and fitness.
Allergische Kontaktdermatitis ist eine zellvermittelte verzögerte Überempfindlichkeitsreaktion auf niedermolekulare Substanzen, die häufig Hautreaktionen hervorrufen und dadurch sowohl persönliche als auch berufliche Probleme verursachen. Chemikalien wie para-Phenylendiamin (PPD) und andere verwandte para-Aminobenzole werden häufig in Farben eingesetzt und erlangen aufgrund ihrer starken Allergenität eine immer größere Aufmerksamkeit. Bis jetzt ist das Wissen über zelluläre Immunantworten auf PPD nur begrenzt verstanden. In dieser Studie haben wir das immunmodulatorische Potential vonrnPPD untersucht, vor allem hinsichtlich der Fragestellung, ob PPD selber naive T-Zellen sensibilisieren kann oder ob PPD letztendlich nur ein Allergen ist. In Anbetracht der Tatsache das dendritische Zellen (DZ) eine vorherrschende Rolle bei der allergischen Kontaktdermatitis spielen, haben wir untersucht, in wie weit PPD in der Lage ist, funktionelle Reifung zu modulieren. Um abschätzen zu können, welcher Anteil an PPD nach der Stimulation zur potentiellen Aktivierung tatsächlich zur Verfügung steht, haben wir die metabolische Kompetenz von "monocyte derived dendritic cells" (MoDC) untersucht, PPD zurnacetylieren. Dazu haben wir die N-Acetyltransferase 1 (NAT-1) und N-Acetyltransferase 2(NAT-2) mRNA Expression charakterisiert und weitergehend die metabolische Aktivität von NAT-1 Enzym bestimmt. Mit diesen Versuchen konnten wir zeigen, dass MoDC von 9 aus 10 Spendern das NAT-1 Enzym exprimieren und konnten in 4 von 6 Spendern acetyliertes PPD in Extrakten aus Zellkulturüberständen detektieren. In Anbetracht der Tatsache, dassrnacetyliertes PPD nicht in der Lage ist, MoDC zu aktivieren, können wir davon ausgehen,dass der Anteil an acetyliertem PPD zur DC Aktivierung nicht zur Verfügung steht. Um das sensibilisierende Potential von PPD herauszufinden, haben wir nach Stimulation mit verschiedenen PPD Konzentrationen die Expression von Oberflächenmolekülen, die für die Antigenpräsentation, Co-Stimulation, späte Migration und dendritische Zell-/T-Zell-Wechselwirkung entscheidend sind, zu verschiedenen Zeitpunkten mittels Durchflusszytometrie Messungen (FACS) gemessen. Zusammengefasst konnten wir zeigen,dass PPD signifikant die "chemokine receptor 7" (CCR7) Expression und nicht signifikant die "human leukocyte antigen " DR" HLA-DR, "DC-specific C-type lectin intercellular adhesion molecule-3-grabbing nonintegrin" (DC-SIGN) und "cluster domain" 11c (CD11c) Expression erhöht. Die erhöhte CCR7 Expression ermutigte uns, weiter nachzuforschen, ob PPD trotz der Tatsache, dass es keine traditionelle CD80 und CD86 Expression induzieren kann, fähig ist, DCs zu aktivieren. Ausgiebige Zytokinkinetikmessungen konnten diese Hypothese bestätigen. Mit Cytometric Beat Array (CBA) Messungen konnten wir gleichzeitig Mediatoren wie "Interleukin -1ß" (IL-1ß), "tumor necrosis factor-alpha" (TNF-α), IL-8, IL-6, IL-12P70 undrnIL-10 messen und damit demonstrieren, dass PPD in der Lage ist, bei allen Spendern "Danger Signals" zu induzieren. Zudem konnten wir zeigen, dass die PPD induzierte Zytokinausschüttung interindividuell schwankte und zu unterschiedlichen Zeitpunkten erfolgte. Überraschenderweise war die qualitative Zusammensetzung der Zytokine vergleichbar. Während in 3 von 4 Spendern die Zytokinexpression eher schwach erhöht war und von kurzer Dauer, haben wir in einem Spender vor allem mit den Konzentrationen von 10μM und 50μM PPD verblüffend hohe Zytokinausschüttungen gemessen. Dieser Spenderrnscheint eine Person zu repräsetieren, die empfänglicher für eine PPD Sensibilisierung ist als andere. Unabhängig davon, hat uns die Tatsache, dass PPD in allen Spendern "Danger Signals", aber keine gesteigerte Expressionsrate von traditionellen Reifungsmarkern (mit Ausnahme von CCR7) induzierte, veranlasst zu untersuchen, ob PPD anderweitig fähig ist,in MoDC Immunantworten zu modulieren. Daraufhin haben wir den Einfluss von PPD aufrnaktivierte DCs gemessen und die Ergebnisse bestätigten die Annahme. Weiterhin konnten wir zeigen, dass PPD immunogenes Potential aufweist. Aufgrund der Tatsache, dass die LPS induzierten Signaltransduktionswege in MoDC gut charakterisiert sind, konnten wir mit dieser Studie zusätzlich Hinweise über die PPD induzierte Signalgebung liefern.
Um die weiterhin ansteigende Weltbevölkerung auch zukünftig mit Nahrungsmitteln und anderen wichtigen Rohstoffen zu versorgen, bedarf es einer immer größeren Nutzungsausweitung der Landwirtschaft auf natürliche Flächen. Im Jahre 2009 wurde ein Drittel der Landoberfläche der Erde bereits landwirtschaftlich genutzt. In jüngster Zeit kommt der Landwirtschaft zusätzlich eine wichtige Bedeutung als Energielieferant zu. Ein weiter steigender Flächenbedarf ist die Konsequenz dieser Entwicklung. Dies führt zum Verlust von Habitaten und somit zu einer starken Fragmentierung der Landschaft. Die direkten und indirekten Auswirkungen dieser Entwicklung auf den weltweiten Biodiversitätsrückgang werden mittlerweile als eine der Hauptursachen für den Rückgang vieler Arten anerkannt. Auch der Gesetzgeber hat dieses Problem erkannt und versucht dem entgegenzuwirken. In der vorliegenden Arbeit wird durch einen multiplen Methodenansatz der Einfluss der Habitatfragmentierung auf die Konnektivität der Populationen von zwei unterschiedlich mobilen Insektenarten untersucht. Als Modelarten wurden eine wenig mobile Heuschreckenart (Chorthippus montanus) und ein hoch mobiler Tagfalter (Brenthis ino) ausgewählt, welche beide an feuchte Grünländer gebunden sind. Für C. montanus wurde gezeigt, dass die Art nur ein sehr eingeschränktes Mobilitätspotenzial besitzt und stark an ihr Habitat gebunden ist. Daher ist eine Durchquerung von ungeeigneten Lebensräumen als äußerst unwahrscheinlich zu erachten. Eine Studie zur Mobilität und dem Verhalten von Brenthis ino belegt eine starke Philopatrie für eine nicht territoriale Insektenart. Zusätzlich zu den Fang-Wiederfang-Untersuchungen im Freiland wurden populations- und landschaftsgenetische Analysen durchgeführt. Die populationsgenetischen Analysen zeigen eine starke Isolation der Populationen von C. montanus, während zwischen den untersuchten Populationen von B. ino eine gute Konnektivität besteht. Mittels der landschaftsgenetischen Analyse wird gezeigt, dass Wälder und bebaute Flächen für beide Arten substanzielle Genefluss-Barrieren darstellen. Basierend auf einer GIS gestützten Analyse wird dargelegt, dass eine lokal ausreichende Vernetzung von Grünländern in Rheinland-Pfalz durch den Biotopverbund existiert. Die Berücksichtigung der Konnektivität bei der Auswahl und Ausgestaltung der Instrumente zur rechtlichen Sicherung und Umsetzung des Biotopverbundes (-§ 21 BNatSchG) bergen großes Potenzial zum Schutz von Grünländern. Ferner wird gezeigt, dass die Landschaftsplanung zwar ein geeignetes Instrument zur Erarbeitung des Biotopverbundes ist, aber kein Instrument zur rechtlichen Sicherung desselbigen im Sinne des -§ 21 Abs. 4 BNatSchG sein kann.
Tropospheric ozone (O3) is known to have various detrimental effects on plants, such as visible leaf injury, reduced growth and premature senescence. Flux models offer the determination of the harmful ozone dose entering the plant through the stomata. This dose can then be related to phytotoxic effects mentioned above to obtain dose-response relationships, which are a helpful tool for the formulation of abatement strategies of ozone precursors. rnOzone flux models are dependant on the correct estimation of stomatal conductance (gs). Based on measurements of gs, an ozone flux model for two white clover clones (Trifolium repens L. cv Regal; NC-S (ozone-sensitive) and NC-R (ozone-resistant)) differing in their sensitivity to ozone was developed with the help of artificial neural networks (ANNs). White clover is an important species of various European grassland communities. The clover plants were exposed to ambient air at three sites in the Trier region (West Germany) during five consecutive growing seasons (1997 to 2001). The response parameters visible leaf injury and biomass ratio of NC-S/NC-R clone were regularly assessed. gs-measurements of both clones functioned as output of the ANN-based gs model, while corresponding climate parameters (i.e. temperature, vapour pressure deficit (VPD) and photosynthetic active radiation (PAR)) and various ozone concentration indices were inputs. The development of the model was documented in detail and various model evaluation techniques (e.g. sensitivity analysis) were applied. The resulting gs model was used as a basis for ozone flux calculations, which were related to above mentioned response parameters. rnThe results showed that the ANNs were capable of revealing and learning the complex relationship between gs and key meteorological parameters and ozone concentration indices. The dose-response relationships between ozone fluxes and visible leaf injury were reasonably strong, while those between ozone fluxes and NC-S/NC-R biomass ratio were fairly weak. The results were discussed in detail with respect to the suitability of the chosen experimental methods and model type.
Comparing the results of the phylogeographies of the four species included in this thesis, some accordances have been found, even though certain patterns are only represented in one or two species. In all cases, the findings of the studied species strongly support the existence of forests or forest-like ecosystems beyond the classic forest refugia in the Mediterranean areas (Iberian, Apennine and Balkan peninsulas) during glacial times. However, evidence of glacial refugial areas in Southeastern Europe, especially the Balkans, have been found in this study as well. The analysed populations of Aposeris foetida, Melampyrum sylvaticum and Erebia euryale showed high genetic diversity values and mostly higher private fragments in this area, which is a strong indicator for centres of glacial survival during Würm and, regarding the results of M. sylvaticum, even during the Riss ice age. Three of the analysed species (A. foetida, M. sylvaticum and Colias palaeno) supported a second main glacial refuge area located along the Northern Alps. Again, high genetic diversity values and the uniqueness of the populations living in this region today prove the importance of this area as a glacial centre of survival. Those results confirm several recently published studies on forest species and strongly indicate the persistence of forest-like structures or even forests during the ice ages along the foothills of the Northern Alps. Additionally, the persistence of C. palaeno in this area furthermore supports the existence of peatlands north of the Alps, at least during the last glacial. The results of M. sylvaticum and E. euryale further indicate the vicinity of the Tatra Mountains as core areas for glacial survival. However, the genetic patterns found for E. euryale are ambiguous. Due to an intermediate position of two genetic lineages (originating in the Eastern Alps and Southeastern Europe), the Tatras could also reflect a postglacial mixture zone of those lineages. Moreover, the glacial and postglacial importance of this area for woodland species was accentuated, supporting other phylogeographic studies published. Besides the congruities among the results of the study species, some unique patterns and therefore further potential glacial refugia have also been illuminated in this thesis. For instance, the calcicole species, A. foetida, most probably had further survival area at both sides of the Dinaric Alps, supported by high genetic diversity values and a high number of private fragments found in Croatian populations. Furthermore, the surroundings of the German Uplands and the margin of the Southern Alps provided suitable conditions for glacial survival for M. sylvaticum, while the Eastern and Southeastern Alpine region most probably sheltered the Large Ringlet E. euryale during ice ages. Additionally, this butterfly species survived at least the glaciation along the foothills of the Massif Central, whose present populations showed a unique genetic lineage and their genetic diversity values have been measurably higher than in other populations for this species. Finally, a large and continuous Würm distribution is highly likely south of the Fennoscandian glaciers in Central Europe for C. palaeno, which might indicate extended peatland areas during Würm glacial. With all the patterns found in this study, the understanding of glacial persistence of forest, respectively forest-like structures and peatlands during Würm or even Riss glacial in Europe could be advanced. The congruencies among the analysed woodland and bog species illustrate the importance and location of extra-Mediterranean refugia for European mountain forests and the glacial presence of Central European peatlands. Thus, already postulated theories could be supported and further pieces of the overall puzzle could be added. The varieties of the different survival centres once more clarified that further phylogeographic studies on mountain forest of different habitat requirements and especially peatland species have to be implemented to get a clearer picture of the glacial history of these habitats.
Climate change and habitat fragmentation modify the natural habitat of many wetland biota and lead to new compositions of biodiversity in these ecosystems. While the direct effects of climate are often well known, indirect effects due to biotic interactions remain poorly understood. The water meadow grasshopper, Chorthippus montanus, is a univoltine habitat specialist, which is adapted to permanently moist habitats. Land use change and drainage led to highly fragmented populations of this generally flightless species. In large parts of the Palaearctic Ch. montanus occurs sympatrically with its widespread congener, the meadow grasshopper Chorthippus parallelus. Due to their close relationship and their similar songs, hybridization is likely to occur in syntopic populations. Such a species pair of a habitat specialist and a habitat generalist represents an ideal model system to examine the role of ongoing climate change and an accumulation of extreme climatic events on the life history strategies, population dynamics and inter-specific interactions. In Chapter I a laboratory experiment was conducted to identify the impact of environmental factors on intra-specific life-history traits of Ch. montanus. Like other Orthoptera species, Ch. montanus follows a converse temperature size rule. In line with the dimorphic niche hypothesis, which states that sexual size dimorphism evolved in response to the different sexual reproductive roles, both sexes showed different responses to increasing density at lower temperatures. Males attained smaller body sizes at high densities, whereas females had a prolonged development time. This is the first evidence for a sex-specific phenotypic plasticity in Ch. montanus. Females benefit from the prolonged development as their reproductive success depends on the size and number of egg clutches they may produce. By contrast, the reproductive success of males depends on the chance to fertilize virgin females, which increases with faster development. This may become a disadvantage for Ch. montanus as an intraspecific phenology shift may increase hybridization risk with the sibling species. Despite the widespread assumption that hybridization between two sympatric species is rare due to complete reproductive barriers, the genetic analyses of 16 populations (Chapter II) provided evidence for wide prevalence of hybridization between both species in the wild. As no complete admixture was found in the examined population, it is assumed that hybridization only occurs in ecotones between wetlands and drier parts. Reproductive barriers (habitat isolation, behavior, phenology) seem to prevent the genetic swamping of Ch. montanus populations. Although a behavioral experiment showed that mate choice presents an important reproductive barrier between both species, the experiment also revealed that reproductive barriers could be altered by environmental change (e.g. increasing heterospecific frequency). Chapter III analyzes the impact of extreme climatic events on population dynamics and interspecific hybridization. A mark-recapture analysis combined with weather records over five years provides evidence that the embryonic development in Ch. montanus is vulnerable to extreme climatic events. Strong population declines in Ch. montanus lead to a disequilibrium between Ch. montanus and Ch. parallelus populations and increases the risk of hybridization. The highest hybridization risk was found in the first weeks of a season, when both species had an overlapping phenology. Furthermore, hybrids were generally localized at the edge of the Ch. montanus distribution with higher heterospecific encounter probabilities. The hybridization rate reached up to 19.6%. The genetic analyses in Chapter II and III show that hybridization differentially affects specialists and generalists. While generalists may benefit from hybridization by an increasing genetic diversity, such a positive correlation was not found for Ch. montanus. The results underline the importance of reproductive barriers for the co-existence of these sympatric species. However, climate change and other anthropogenic disturbances alter reproductive barriers and promote hybridization, which may threaten small populations by genetic displacement. As anthropogenic hybridization is recognized as a major threat to biodiversity, it should be considered in environmental law and policy. In Chapter IV the role of hybrids and hybridization in three levels of law and the historical backgrounds of hybrids becoming a part of legal instruments is analyzed. Due to legal uncertainties and the complexity of this topic a legal assessment of hybrids is challenging and argues for species-specific approaches. Nonetheless, existing legal norms provide a suitable basis, but need to be specified. Finally, this chapter discusses different opportunities for the management of hybrids and hybridization in a conservation perspective and their necessity.
N-acetylation by N-acetyltransferase 1 (NAT1) is an important biotransformation pathway of the human skin and it is involved in the deactivation of the arylamine and well-known contact allergen para-phenylenediamine (PPD). Here, NAT1 expression and activity were analyzed in antigen presenting cells (monocyte-derived dendritic cells, MoDCs, a model for epidermal Langerhans cells) and human keratinocytes. The latter were used to study exogenous and endogenous NAT1 activity modulations. Within this thesis, MoDCs were found to express metabolically active NAT1. Activities were between 23.4 and 26.6 nmol/mg/min and thus comparable to peripheral blood mononuclear cells. These data suggest that epidermal Langerhans cells contribute to the cutaneous N-acetylation capacity. Keratinocytes, which are known for their efficient N-acetylation, were analyzed in a comparative study using primary keratinocytes (NHEK) and different shipments of the immortalized keratinocyte cell line HaCaT, in order to investigate the ability of the cell line to model epidermal biotransformation. N-acetylation of the substrate para-aminobenzoic acid (PABA) was 3.4-fold higher in HaCaT compared to NHEK and varied between the HaCaT shipments (range 12.0"44.5 nmol/mg/min). Since B[a]P induced cytochrome p450 1 (CYP1) activities were also higher in HaCaT compared to NHEK, the cell line can be considered as an in vitro tool to qualitatively model epidermal metabolism, regarding NAT1 and CYP1. The HaCaT shipment with the highest NAT1 activity showed only minimal reduction of cell viability after treatment with PPD and was subsequently used to study interactions between NAT1 and PPD in keratinocytes. Treatment with PPD induced expression of cyclooxygenases (COX) in HaCaT, but in parallel, PPD N-acetylation was found to saturate with increasing PPD concentration. This saturation explains the presence of the PPD induced COX induction despite the high N-acetylation capacities. A detailed analysis of the effect of PPD on NAT1 revealed that the saturation of PPD N-acetylation was caused by a PPD-induced decrease of NAT1 activity. This inhibition was found in HaCaT as well as in primary keratinocytes after treatment with PPD and PABA. Regarding the mechanism, reduced NAT1 protein level and unaffected NAT1 mRNA expression after PPD treatment adduced clear evidences for substrate-dependent NAT1 downregulation. These results expand the existing knowledge about substrate-dependent NAT1 downregulation to human epithelial skin cells and demonstrate that NAT1 activity in keratinocytes can be modulated by exogenous factors. Further analysis of HaCaT cells from different shipments revealed an accelerated progression through the cell cycle in HaCaT cells with high NAT1 activities. These findings suggest an association between NAT1 and proliferation in keratinocytes as it has been proposed earlier for tumor cells. In conclusion, N-acetylation capacity of MoDCs as well as keratinocytes contribute to the overall N-acetylation capacity of human skin. NAT1 activity of keratinocytes and consequently the detoxification capacities of human skin can be modulated by the presence of exogenous NAT1 substrates and endogenous by the cell proliferation status of keratinocytes.