Academic literature on the topic 'Temperature-dependent sex determination'

Abstract There are a remarkable variety of sex determination systems among different animal taxa. In most animals, sex is determined chromosomally. Although in an increasing number of animals sex determination has been found to be influenced primarily by the environment. Species with genotypic sex determination (GSD) have their sex determined at the time of fertilization, by genetic factors alone and those with environmental sex determination (ESD) have their sex determined by environmental factors that act after fertilization. Temperature-dependent Sex Determination (TSD), whereby the sex of the developing embryos depends on the temperature at which they develop is widespread in oviparous reptiles and occurs in all crocodilians, marine turtles and tuatara examined to date and is common in many freshwater turtles and lizards. SECTION ONE Temperature-dependent sex determination (TSD) was never expected to occur in viviparous reptiles, as thermoregulation by pregnant females would result in relatively stable gestation temperatures. Temperature-dependent sex determination and viviparity goes against all the basic assumptions that TSD occurs in oviparous reptiles where temperatures within a nest vary widely. However, skewed sex ratios as a result of incubation temperature indicated the possibility of TSD in the viviparous lizard Eulamprus tympanum. In my first experiments I show the first recorded case of a viviparous reptile with TSD. The developing embryos of the viviparous skink E. tympanum are subject to TSD, with gestation temperature having a highly significant effect on sex and warmer temperatures giving rise to male offspring (Chapter 1). Sex is fully determined at the time of birth and can be differentiated histologically into testes or ovaries (Chapter 2). The morphology and histological characteristics of the gonads of neonatal E. tympanum resulting from the treatment temperatures described in chapter 1 illustrate that sex in E. tympanum is easily distinguished at the time of birth and corresponds with the presence or absence of hemipenes. Males are histologically characterised by an elongated gonad consisting of seminiferous tubules with either no cortical epithelium or, if present at all, in a very thin band. If they are present, Mϋllerian ducts, showing signs of degeneration, are attached to the kidney by a shortened mesosalpinx. Females are histologically characterised by an irregularly shaped gonad consisting of a thick cortical epithelium that occasionally contains oocytes. The Mϋllerian ducts are obvious structures attached to the kidney by a fibrous mesosalpinx. The presence or absence of hemipenes is a reliable technique for determining sex in newborn E. tympanum. Sex determination is easiest to perform on neonates within the first few days of birth as hemipenes become increasingly difficult to evert as neonates age, however, with practice they are easily identified without full eversion. SECTION TWO The thermal biology of E. tympanum in the field is restricted by both the thermal properties of their habitat (Chapter 3) and behavioural modifications when faced with a predation threat (Chapter 4). The available temperatures in the field suggest that TSD is biologically relevant in the species and not just a laboratory artefact; E. tympanum can attain mean selected temperatures achieved in the laboratory but the proportion of time at the temperature is restricted. Females actively thermoregulate in the field, although they are restricted in their efficiency of thermoregulation by environmental constraints, for example, microhabitat structure, weather conditions, predator avoidance and social ranking. The highly territorial nature and high densities of E. tympanum present in Kanangra Boyd National Park potentially force less dominant individuals into less favourable habitats that are significantly cooler. An important point is that gravid females in more favourable habitats in the period encompassing the middle third of development (the assumed sex determining period) are selecting higher temperatures, with lower variance and have greater thermoregulatory efficiency than during the rest of pregnancy, therefore, thermoregulating more precisely during this thermosensitive period (Chapter 3). Chemosensory cues provide important information on the risk of predation. Hence, chemoreception is a common mechanism used by many species to detect the presence of, and subsequently respond to, a potential predator. The perceived risk of predation may force retreat to sub-optimal conditions, forcing a trade-off between the risk of predation and the ability to acquire resources. The basking regime maintained by gravid female E. tympanum, can directly alter sex ratios of offspring produced through temperature-dependent sex determination (Chapter 1). The avoidance of predator scents may restrict basking ability and in turn alter the sex of offspring produced. I measured responsiveness to chemical cues using tongue flicks as an indicator of chemical discrimination in females of different reproductive condition. I then measured activity and basking behaviour of gravid and non-gravid females in experimental enclosures in the presence of various chemical stimuli to determine if basking opportunity is compromised by the presence of a predator scent. Females respond differently depending upon reproductive condition, with gravid females responding most significantly to a predator scent. Activity, basking frequency, and time spent in the open (basking duration) are significantly reduced in gravid females in the presence of a predator stimulus. Under laboratory conditions, gravid females modify their behaviour and forego the opportunity to bask when there is a perceived predation risk (Chapter 4). SECTION THREE As female viviparous reptiles can regulate the temperature of the embryo by maternal temperature selection (Chapter 1), the occurrence of TSD in E. tympanum opens the possibility for females to select the sex of offspring. Reproducing females may benefit by facultatively adjusting their investment into sons over daughters or vice versa, in response to population wide shifts in adult sex ratios. Female E. tympanum, can manipulate the sex of their offspring in response to sex imbalances in the population using temperature-dependent sex determination (Chapter 5). When adult males are scarce, females produce male-biased litters and when adult males are common, females produce female-biased litters. The cues used by a female to assess the adult population are not known, but presumably depends upon the female's experience throughout the breeding season and is the subject of further investigation (Chapter 6). The maternal manipulation of offspring sex ratio in E. tympanum suggests a selective advantage of temperature-dependent sex determination. Any facultative sex ratio response needs to recognise the scarcity of one sex in order to overproduce that sex in the next generation; offspring sex ratio will vary inversely with adult sex ratio. Maternal sex allocation in E. tympanum is linked with population (or adult) sex ratio (Chapter 5), and one of the mechanisms by which females recognise an imbalance may be linked to visual recognition of males (Chapter 6). Females maintained throughout pregnancy without any male stimulus produce entirely male offspring (Chapter 5). In contrast females exposed to male stimulus produce both sexes (Chapter 5). Females respond differently to varying degrees of male stimulus and visual recognition of males in a population may be more important than chemoreception. In the absence of visual cues, females produce more male offspring, even when chemosensory cues are present (Chapter 6). The study system presented here offers many advantages over oviparous species with TSD, due to E. tympanum being relatively short lived and fast maturing. Thus, the fitness consequences over multiple generations as a result of gestation can be investigated. Viviparity allows maternal control of embryonic temperature during gestation and a means of maternal sex allocation. Until now the maternal side of TSD and sex allocation has been where the mother deposits her eggs and the allocation of sex steroid hormones at oviposition, both of which have been difficult to study. The work presented and the study system itself should inspire great interest in TSD and viviparous reptiles.

Abstract There are a remarkable variety of sex determination systems among different animal taxa. In most animals, sex is determined chromosomally. Although in an increasing number of animals sex determination has been found to be influenced primarily by the environment. Species with genotypic sex determination (GSD) have their sex determined at the time of fertilization, by genetic factors alone and those with environmental sex determination (ESD) have their sex determined by environmental factors that act after fertilization. Temperature-dependent Sex Determination (TSD), whereby the sex of the developing embryos depends on the temperature at which they develop is widespread in oviparous reptiles and occurs in all crocodilians, marine turtles and tuatara examined to date and is common in many freshwater turtles and lizards. SECTION ONE Temperature-dependent sex determination (TSD) was never expected to occur in viviparous reptiles, as thermoregulation by pregnant females would result in relatively stable gestation temperatures. Temperature-dependent sex determination and viviparity goes against all the basic assumptions that TSD occurs in oviparous reptiles where temperatures within a nest vary widely. However, skewed sex ratios as a result of incubation temperature indicated the possibility of TSD in the viviparous lizard Eulamprus tympanum. In my first experiments I show the first recorded case of a viviparous reptile with TSD. The developing embryos of the viviparous skink E. tympanum are subject to TSD, with gestation temperature having a highly significant effect on sex and warmer temperatures giving rise to male offspring (Chapter 1). Sex is fully determined at the time of birth and can be differentiated histologically into testes or ovaries (Chapter 2). The morphology and histological characteristics of the gonads of neonatal E. tympanum resulting from the treatment temperatures described in chapter 1 illustrate that sex in E. tympanum is easily distinguished at the time of birth and corresponds with the presence or absence of hemipenes. Males are histologically characterised by an elongated gonad consisting of seminiferous tubules with either no cortical epithelium or, if present at all, in a very thin band. If they are present, M�llerian ducts, showing signs of degeneration, are attached to the kidney by a shortened mesosalpinx. Females are histologically characterised by an irregularly shaped gonad consisting of a thick cortical epithelium that occasionally contains oocytes. The M�llerian ducts are obvious structures attached to the kidney by a fibrous mesosalpinx. The presence or absence of hemipenes is a reliable technique for determining sex in newborn E. tympanum. Sex determination is easiest to perform on neonates within the first few days of birth as hemipenes become increasingly difficult to evert as neonates age, however, with practice they are easily identified without full eversion. SECTION TWO The thermal biology of E. tympanum in the field is restricted by both the thermal properties of their habitat (Chapter 3) and behavioural modifications when faced with a predation threat (Chapter 4). The available temperatures in the field suggest that TSD is biologically relevant in the species and not just a laboratory artefact; E. tympanum can attain mean selected temperatures achieved in the laboratory but the proportion of time at the temperature is restricted. Females actively thermoregulate in the field, although they are restricted in their efficiency of thermoregulation by environmental constraints, for example, microhabitat structure, weather conditions, predator avoidance and social ranking. The highly territorial nature and high densities of E. tympanum present in Kanangra Boyd National Park potentially force less dominant individuals into less favourable habitats that are significantly cooler. An important point is that gravid females in more favourable habitats in the period encompassing the middle third of development (the assumed sex determining period) are selecting higher temperatures, with lower variance and have greater thermoregulatory efficiency than during the rest of pregnancy, therefore, thermoregulating more precisely during this thermosensitive period (Chapter 3). Chemosensory cues provide important information on the risk of predation. Hence, chemoreception is a common mechanism used by many species to detect the presence of, and subsequently respond to, a potential predator. The perceived risk of predation may force retreat to sub-optimal conditions, forcing a trade-off between the risk of predation and the ability to acquire resources. The basking regime maintained by gravid female E. tympanum, can directly alter sex ratios of offspring produced through temperature-dependent sex determination (Chapter 1). The avoidance of predator scents may restrict basking ability and in turn alter the sex of offspring produced. I measured responsiveness to chemical cues using tongue flicks as an indicator of chemical discrimination in females of different reproductive condition. I then measured activity and basking behaviour of gravid and non-gravid females in experimental enclosures in the presence of various chemical stimuli to determine if basking opportunity is compromised by the presence of a predator scent. Females respond differently depending upon reproductive condition, with gravid females responding most significantly to a predator scent. Activity, basking frequency, and time spent in the open (basking duration) are significantly reduced in gravid females in the presence of a predator stimulus. Under laboratory conditions, gravid females modify their behaviour and forego the opportunity to bask when there is a perceived predation risk (Chapter 4). SECTION THREE As female viviparous reptiles can regulate the temperature of the embryo by maternal temperature selection (Chapter 1), the occurrence of TSD in E. tympanum opens the possibility for females to select the sex of offspring. Reproducing females may benefit by facultatively adjusting their investment into sons over daughters or vice versa, in response to population wide shifts in adult sex ratios. Female E. tympanum, can manipulate the sex of their offspring in response to sex imbalances in the population using temperature-dependent sex determination (Chapter 5). When adult males are scarce, females produce male-biased litters and when adult males are common, females produce female-biased litters. The cues used by a female to assess the adult population are not known, but presumably depends upon the female�s experience throughout the breeding season and is the subject of further investigation (Chapter 6). The maternal manipulation of offspring sex ratio in E. tympanum suggests a selective advantage of temperature-dependent sex determination. Any facultative sex ratio response needs to recognise the scarcity of one sex in order to overproduce that sex in the next generation; offspring sex ratio will vary inversely with adult sex ratio. Maternal sex allocation in E. tympanum is linked with population (or adult) sex ratio (Chapter 5), and one of the mechanisms by which females recognise an imbalance may be linked to visual recognition of males (Chapter 6). Females maintained throughout pregnancy without any male stimulus produce entirely male offspring (Chapter 5). In contrast females exposed to male stimulus produce both sexes (Chapter 5). Females respond differently to varying degrees of male stimulus and visual recognition of males in a population may be more important than chemoreception. In the absence of visual cues, females produce more male offspring, even when chemosensory cues are present (Chapter 6). The study system presented here offers many advantages over oviparous species with TSD, due to E. tympanum being relatively short lived and fast maturing. Thus, the fitness consequences over multiple generations as a result of gestation can be investigated. Viviparity allows maternal control of embryonic temperature during gestation and a means of maternal sex allocation. Until now the maternal side of TSD and sex allocation has been where the mother deposits her eggs and the allocation of sex steroid hormones at oviposition, both of which have been difficult to study. The work presented and the study system itself should inspire great interest in TSD and viviparous reptiles.

Thesis (Ph. D.)--University of Alabama at Birmingham, 2007.
Additional advisors: Asim Bej, Gene Hines, Douglas Watson, Douglas Weigent. Description based on contents viewed Oct. 2, 2008; title from PDF t.p. Includes bibliographical references.

Thesis (Ph. D.)--University of Sydney, 2004.
Title from title screen (viewed 5 May 2008). Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy to the School of Biological Sciences, Faculty of Science. Degree awarded 2004; thesis submitted 2003. Appendices contains published articles co-authored by Robert. Includes bibliographical references. Also available in print form.

Captive husbandry programs in zoos have documented nesting behavior and have successfully hatched Manouria emys emys, but data on sex determining mechanisms and sex ratios are absent. A total of 30 M. e. emys eggs were artificially incubated at five different temperatures in constant humidity. Mean incubator temperatures were 24.99°C, 25.06°C, 27.18°C, 28.00°C, and 30.79°C. Incubation duration ranged from 60 days to 92 days, and hatching success was 50%. Sex determined by histology and laparoscopy resulted in male differentiation at low temperatures (24.99°C, 27.18°C) and female differentiation at high temperatures (30.79°C). Pivotal temperature was estimated to be 29.29°C. The following investigation into temperature-dependent sex determination (TSD), including its presence or absence, pattern, and pivotal temperature, has implications for studies of adaptive significance of reproductive behaviors and of chelonian phylogenetic history. Additionally, the proposed study can provide foundations for conservation management decisions, and for captive breeding programs.

In many reptile species, offspring sex is determined by the temperature these animals experience during embryogenesis, rather than by genetic factors passed from parents to offspring (e.g., sex chromosomes). The adaptive value of this unusual sex-determining mechanism (temperature-dependent sex determination, TSD) has eluded satisfactory explanation since its discovery four decades ago. The most plausible suggestion in this regard (the Charnov-Bull model) proposes that TSD enhances maternal fitness when nest temperature has a differential impact on the fitness of sons versus daughters, such that male-producing temperatures are optimal for the fitness of sons, and female-producing temperatures are optimal for the fitness of daughters. However, because most reptiles with TSD are long-lived and have delayed sexual maturation, no robust experimental test of this model has been conducted. The primary goal of my PhD research was to experimentally test the Chamov-Bull model using a_ short—lived, early-maturing lizard with TSD (the jacky dragon, Amphibolurus muricatus). By incubating eggs at a range of temperatures, and using hormonal manipulations to de-confound the effects of incubation temperature and sex on offspring fitness, my results directly address (and strongly support) major predictions of the Charnov—Bull model. Overall, males from eggs that were incubated at male-producing temperatures were more successful at siring offspring than those from eggs incubated under female-producing temperatures. This pattern was reversed for females; female reproductive success was lowest for individuals incubated at male-producing temperatures. In addition, incubation temperature had a strong effect on the seasonal timing of hatching, and the optimal time of hatching is likely to differ between sons versus daughters. Hatching early in the season provided individuals with a relatively long growing season, thereby enabling offspring to reach sexual maturity by age one. In A. muricatus, the fitness difference between early—maturing sons versus daughters depends upon the intensity of competition for mating opportunities. Early-maturing sons are unlikely to reproduce in their first year because of intense competition with larger territorial individuals from previous cohorts, whereas older females do not suppress breeding of younger females in the same way. These patterns suggest that a daughter's (but not a son’s) reproductive success would be enhanced if she hatched early enough to reach sexual maturity by age one. Thus, TSD should enhance maternal fitness by enabling the overproduction of daughters early in the season, and males late in the season. My second objective was to evaluate sex allocation patterns in A. muricatus, and especially to clarify the effects of maternal factors on offspring sex ratios. By rearing reproductive females and eggs under standardized conditions, my results suggested a strong maternal (perhaps genetic) component to sex determination. Clutch sex ratios varied substantially among females, and the degree of this variation depended upon the timing of clutch production. Moreover, this variation in clutch sex ratios (as well as other offspring phenotypes) was not associated with any non—genetic maternal effects, such as egg size, yolk steroid allocation, or nest-site selection. However, additional manipulative studies demonstrated that maternal sex allocation was responsive to the quality of the diet provided to reproducing females, as well as the operational sex ratios experienced prior to and during the reproductive season. Overall, these results challenge current paradigms of reptilian TSD, suggesting that offspring sex is likely the result of complex interactions among multiple factors, rather than that of a single overriding variable (i.e., temperature). My third objective was to complement my experimental studies with field-based observations, to provide insight into the ecological relevance of my laboratory-based results. A mark-recapture study in the field demonstrated that early hatching enhances offspring fitness in numerous ways. For example, individuals that hatched early in the season grew faster, dispersed farther, and had greater survival rates than those that hatched late. Moreover, early-hatched individuals attained sexual maturity by age one under natural field conditions, strongly supporting findings from my experimental work. I used radiotelemetry to track free-ranging gravid females and locate their nest sites, enabling me to evaluate nesting behaviour, as well as effects of natural nest conditions on offspring phenotypes and sex ratios. Females selected nest sites with lower canopy cover, and hence higher temperatures, than random. Moreover, seasonal shifts in ambient temperatures caused concomitant seasonal increases in nest temperature, suggesting that daughters will be over—produced early in the season and males produced later, as predicted by theory. Surprisingly, however, this expectation was not met: offspring sex ratios were not significantly associated with mean nest temperatures (but were linked to the mean daily thermal range). Overall, the field data corroborate many results from my experimental - studies, but suggests a hitherto-unrecognised complexity in the pathways by which sex is determined, and in the ways in which nest temperature affects offspring sex.

Reptiles exhibit marked diversity in sex-determining mechanisms. Many species exhibit genotypic sex determination (GSD) with male heterogamety (XX females/XY males), others have GSD with female heterogamety (ZW females/ZZ males), and still others exhibit temperature-dependent sex determination (TSD). The distribution of these mechanisms throughout the reptile phylogeny implies evolutionary lability in sex determination, and in some lineages there has been a number of transitions between GSD and TSD. Despite this diversity, GSD and TSD have traditionally been viewed as mutually-exclusive mechanisms of sex determination in reptiles, since there is little evidence for their co-occurrence. Considerable empirical and theoretical effort has been directed towards understanding the adaptive significance of TSD in reptiles. In comparison, there has been little focus on understanding how evolutionary transitions between GSD and TSD occur at a genetic and mechanistic level. I addressed this question by applying both empirical and theoretical approaches to investigate interaction of genotypic and temperature influences in the sex determination of two endemic species of Australian lizards. The three-lined skink, Bassiana duperreyi, has XX/XY chromosomal sex determination, yet a previous investigation reported a significant male bias in the sex ratio of eggs incubated at low temperatures. To enable an explicit test for temperature induced sex reversal in this species, a 185 bp Y chromosome marker was isolated by Amplified Fragment Length Polymorphism (AFLP) analysis. The marker was subsequently converted into a duplex PCR assay that co-amplified a 185 bp (or 92 bp) Y chromosome fragment and a 356 bp fragment of the single-copy nuclear gene C-mos (from both sexes) as a positive control. The accuracy of the PCR sex assay was tested on 78 individuals for which sex reversal was not expected. PCR genotype and sex phenotype were concordant for 96% of the animals. This is one of the very few sex tests developed for a reptile, and the first report of Y chromosome sequence from a reptile. The PCR assay was subsequently applied to genotype hatchlings from both cool (16-7.5C) and warm (22-7.5C) cyclical incubation temperature treatments, and identified sex reversal in 15% of genotypically female (XX) embryos (n=26) from the cool treatment, but no sex reversal in eggs from the warmer treatment (n=35). Thus, low incubation temperatures can over-ride genotypic sex determination in B. duperreyi, indicating that GSD and TSD co-occur in this species. The Central bearded dragon, Pogona vitticeps (Agamidae), has ZZ/ZW chromosomal sex determination, and is a member of a lizard family in which GSD and TSD are both widespread, indicating evolutionary lability in sex determination. AFLP analysis was applied to isolate homologous Z and W chromosome-linked markers (71 bp and 72 bp, respectively) from this species. The AFLP sequences were subsequently extended into larger genomic fragments by a reiterated genome walking procedure, producing three non-overlapping contigs of 1.7 kb, 2.2 kb and 4.5 kb. The latter two fragments were verified as distinct, homologous Z/W chromosome fragments by PCR analyses. An amplified 3 kb fragment of the 4.5 kb contig was physically mapped to metaphase spreads, identifying the W microchromosome, and for the first time in this species, the Z microchromosome. PCR analyses indicated the presence of homologous sequences in other Australian agamid species, including both GSD and TSD species. The isolated sequences should therefore prove useful as a comparative genomic tool for investigating the genomic changes that have occurred in evolutionary transitions between sexdetermining mechanisms in agamids, by enabling the identification of chromosomes in TSD species that are homologous to the sex chromosomes of P. vitticeps. The isolated sequences were further converted into a duplex DNA sex assay that co-amplified a 224 bp W chromosome fragment and a 963 bp positive control fragment in both sexes. This PCR assay diagnosed chromosomal sex in three Pogona species, but was not effective outside the genus. Incubation treatment of P. vitticeps eggs revealed a strong and increasing female bias at high constant temperatures (34-36C), but an unbiased sex ratio between 22-32C. Hatchlings from three clutches split between 28C and 34 or 36C incubation treatments were genotyped with the W chromosome AFLP marker. At 28C, the sex ratio was 1:1 but the high temperature treatments produced 2 males and 33 females. All but one of the 30 lizards (97%) incubated at 28C had concordant sex phenotype and genotype, but only 18 of 35 animals (51%) from the high temperature treatment were concordant. All discordant animals were genotypic males (ZZ) that developed as females. Thus, temperature and genotypic influences can interact to determine sex in P. vitticeps. These empirical findings for B. duperreyi and P. vitticeps were extended into a novel theory for the evolution of sex-determining mechanisms in reptiles, working within the framework that species with temperature-induced reversal of chromosomal sex determination are a window to transitional stages of evolution between GSD and TSD. A model was derived from the observation that in both lizards, an extreme of incubation temperature causes sex reversal of the homogametic genotype. In this model, the strength of a genetic regulatory signal for sex determination must exceed a threshold for development of the homogametic sex to occur (male in Pogona, female in Bassiana). The strength of this signal is also temperature-sensitive, so diminishes at extremes of temperature. Simulation modelling demonstrated that increasing the relative magnitude of the threshold for sexual development can cause evolutionary transitions between GSD and TSD. Even more remarkably, decreasing the relative magnitude of the threshold value causes an evolutionary transition between female and male heterogametic GSD. Quantitative adjustment of a single model parameter (the threshold value) thus charts a continuous evolutionary pathway between the three principal mechanisms of sex determination in reptiles (XX/XY-ZZ/ZW-TSD), which were previously considered to be qualitatively distinct mechanisms. The experimental demonstration of temperature-induced reversal of chromosomal sex determination in both B. duperreyi and P. vitticeps presents a challenge to the traditional view that reptilian sex determination is strictly dichotomous (GSD or TSD), and suggests instead that sex determination in reptiles consists of a continuum of systems of interaction between genotypic and temperature influences. Simulation modelling provided solid theoretical support for this proposition, demonstrating that transitions along this continuum are effected simply through shifts in the mean population value for the sex-determining threshold, without requiring substantial genotypic innovation. An important implication of this theory is that transitions between XX/XY and ZZ/ZW modes of GSD may retain the same sex chromosome pair, and the same primary sexdetermining gene, in contrast to previous models for heterogametic transitions. A more immediate implication of these findings is that many reptile species believed to have strict TSD (in particular, lizards and crocodilians), may in fact have a sex-determining system of GSD-TSD interaction, where there is an equilibrium between GSD and TSD individuals within the population.

The prediction and the systematic suppression of self-sustained combustion instabilities in combustors for gas turbine applications still suffer from incomplete physical understanding of the feedback mechanisms and lack of experimental data of the dynamic flame characteristics of Lean-Premixed swirl flames. Hence, the experimental determination of the flame transfer functions of LP swirl flames was achieved using a mixing unit to generate a time-independent and spatial homogeneous mixture of natural gas and combustion air at the burner exit. The determined LP flame dynamics are strongly affected by the formation and in-phase reaction of coherent vortex structures, well known as drivers of combustion instabilities, that have been visualized with an phase-correlated imaging technique. The results discussed in this paper lead to a basic understanding of the frequency-dependent dynamics of LP swirl flames on periodic disturbances and especially, of the influence of the preheating temperature and the air equivalence ratio on the amplitude responses and phase angle functions. Based on the measurements and theoretical considerations concerning the burning velocity of steady-state premixed flames a physical model and — derived from it — scaling laws for the prediction of unstable operation modes in dependence of main operation parameters of the flame were formulated and validated by measurements.

Abstract Current ASME Section III, Division 5 code provides elastic, simplified inelastic and inelastic analysis options for designing nuclear power plant components for elevated temperature service. These analyses methods may fail to capture the complex creep-fatigue response and damage accumulation in materials at elevated temperatures. Hence, for analysis and design of the nuclear power plant components at elevated temperature, a full inelastic analysis that can simulate creep-fatigue responses may be needed. Existing ASME code neither provides guidelines for using full inelastic analysis nor recommends the type of constitutive model to be used. Hence, a unified rate-dependent constitutive model incorporating a damage parameter will be developed, and its parameters for base metal will be determined. In addition, a full inelastic analysis methodology using this model to analyze the creep-fatigue performance of components for nuclear power applications will be developed. Base metal 800H (BM800H) data are collected from literature to determine constitutive material model parameters. The parameter determination methodology for a constitutive model is discussed. The optimized parameter set for BM 800H at different temperatures will be presented in the paper. Recommendations are provided on the constitutive model selection and its parameter determination techniques. In the future, this work will be continued for diffusion bonded Alloy 800H (DB800H) material, and obtained parameters will be compared.

"Lithium-ion batteries currently used in electric vehicles have strong thermal limitations. Thus, both excessively high and low temperatures lead to an accelerated aging process in form of capacity losses through SEI-layer buildup on the one hand and lithium-plating on the other hand. Additionally, temperature gradients within a cell, as well as temperature differences between individual cells have to be minimized in order to increase the system’s durability. With a properly designed battery-thermal-management (BTM), not only the aging process can be minimized but also extreme cases like thermal runaways can be prevented, which otherwise can cause fatal ecological and economical damage or in the worst-case-scenario exposes and harms vehicle occupants to fire, explosion or intoxication due to a damaged battery cell system. For the application of an efficient BTM, a detailed characterization of the battery’s thermal behavior during the charge and discharge cycle with different loads is essential. This paper deals with high-performance pouch cells, as they have a favorable ratio of heat transfer surface area to battery volume for BTM. However, they also present a particular challenge for the acquisition of the surface temperature distribution, as they do not have a solid shell. An external force must be applied to this cell type to prevent it from inflating, when it is under higher loading conditions, because this volume change is expected to change the cell’s thermal behavior. Therefore, the determination of the surface temperature in the present study was carried out while the cell was fixated and forced to stay in its geometric shape. The mechanical fixation has an influence in the thermal behavior and therefore the temperature distribution should be probed in this configuration but this also makes the optical measurement of temperature more difficult as pressure transmission plates must be made transparent. The measured surface temperature distribution can then be used as an input for a FEM-simulation to provide information on the whole cell’s internal temperature field. To the authors´ knowledge, no thermal investigations in this configuration have been reported. Two different optical temperature measurement methods are applied. The first technique is IR-thermography, which is based on the measurement of heat radiation from the surface, and which is restricted to regions on the surface using small IR-transmitting sapphire windows. The second technique, phosphor thermometry, exploits the temperature dependent luminescence properties of thermographic phosphor materials. The phosphors are applied as a thin coating on the surface of the battery and illuminated by a light source. The resulting luminescence emission in the visible range is detected to infer the temperature of the whole surface, since normal glass can be used. The surface temperature distribution obtained by these two techniques are provided and the advantages or disadvantages of each technique are described in terms of precision, accuracy, sensitivity, temporal resolution, spatial resolution and also implementation expenses. The internal temperature field provided by the heat transfer model for a known surface distribution is also presented and the impact of this integral thermal characterization on the BTM is discussed."

The performance of solid oxide fuel cells is affected by various polarization losses, usually grouped in ohmic, activation and concentration polarization. Under typical operating conditions, these polarization losses are largely dependent on cell materials, electrode microstructures, and cell geometry: as an example, the performance of a tubular cell is strongly limited by the ohmic polarization due to the long current path of electrons, while in a planar cell each of these losses has a comparable effect. It is therefore of interest, in case of planar geometry, to investigate the performance limiting factors. In this paper, a performance evaluation of planar circular-shaped seal-less SOFC cells from InDEC® was performed, with an outline of the limiting factors at reduced temperature. Two different designs of planar cells are considered: both have porous NiO-YSZ anode as mechanical support, NiO-YSZ anode active layer, yttria-stabilized zirconia (YSZ) electrolyte, and only differ for the cathode design: (1) strontium doped lanthanum manganate (LSM)-YSZ cathode functional layer (CFL) and LSM cathode current collector layer (CCCL); (2) yttria doped ceria (YDC) blocking layer and lanthanum strontium cobalt ferrite oxide (LSCF) functional layer. The characterization was performed by taking V-I measurements over a range of temperatures between 650°C and 840°C with hydrogen as fuel, and air as oxidant. The dependence of the cell performance on the various polarization contributions was rationalized on the basis of a analytical model, through a parameter estimation on the experimental data, devoted to the determination of the temperature dependence of the area specific resistance (ASR) and of the cathode exchange current density: in particular, the performance limitation at low temperature is due to activation polarization for ASC1 and ohmic polarization for ASC2. Based on the results of the investigation, it is concluded that LSCF cathodes are really effective for decreasing the cathode activation polarization, allowing the reduction of operating temperature. Finally, a microstructural analysis with SEM and optical microscopy has been performed on the ASC2 cell after the polarization testing. The aim of this investigation was in particular to evaluate the degradation phenomena occurring in the anodic structure and over the interfaces between the various active layers. The ASC2 elastic modulus has also been estimated before and after polarization testing in order to evaluate the decreasing of the mechanical strength of the cell after a complete thermal cycle. The results describe a mechanical degradation of the structure and of the distribution of the phases.

Dynamic mechanical properties are critical in the evaluation of materials with viscoelastic behavior. Various techniques, including dynamic mechanical analysis (DMA), rheology, nanoindentation, and others have been developed for this purpose and typically report complex modulus. Each of these techniques has strengths and weaknesses depending on sample geometry and length scale, mechanical properties, and skill of the user. In many industry applications, techniques may also be blindly applied according to a standard procedure without optimization for a specific sample. This can pose challenges for correct characterization of novel materials, and some techniques are more robust to agnostic application than others. A relative assessment of dynamic mechanical techniques is important when considering the appropriate technique to use to characterize a material. It also has bearing on organizations with limited resources that must strategically select one or two capabilities to meet as broad a set of materials as possible. The purpose of this study was to evaluate the measurement characteristics (e.g., precision and bias) of a selection of six dynamic mechanical test methods on a range of polymeric materials. Such a comprehensive comparison of dynamic mechanical testing methods was not identified in the literature. We also considered other technical characteristics of the techniques that influence their usability and strategic value to a laboratory and introduce a novel use of the House of Quality method to systematically compare measurement techniques. The selected methods spanned a range of length scales, frequency ranges, and prevalence of use. DMA, rheology, and oscillatory loading using a servohydraulic tensile tester were evaluated as traditional bulk techniques. Determination of complex modulus by beam vibration was also considered as a bulk technique. At a small length scale, both an oscillatory nanoindentation method and AFM were evaluated. Each method was employed to evaluate samples of polycarbonate, polypropylene, amorphous PET, and semi-crystalline PET. A measurement systems analysis (MSA) based on the ANOVA methods outlined in ASTM E2782 was conducted using storage modulus data obtained at 1 Hz. Additional correlations over a range of frequencies were tested between rheology/DMA and the remaining methods. Note that no attempts were made to optimize data collection for the test specimens. Rather, typical test methods were applied in order to simulate the type of results that would be expected in typical industrial characterization of materials. Data indicated low levels of repeatability error (<5%) for DMA, rheology, and nanoindentation. Biases were material dependent, indicating nonlinearity in the measurement systems. Nanoindentation and AFM results differed from the other techniques for PET samples, where anisotropy is believed to have affected in-plane versus out-of-plane measurements. Tensile-tester based results were generally poor and were determined to be related to the controllability of the actuator relative to the size of test specimens. The vibrations-based test method showed good agreement with time-temperature superposition determined properties from DMA. This result is particularly interesting since the vibrations technique directly accesses higher frequency responses and does not rely on time-temperature superposition, which is not suitable for all materials. MSA results were subsequently evaluated along with other technical attributes of the instruments using the House of Quality method. Technical attributes were weighted against a set of “user demands” that reflect the qualitative expectations often placed on measurement systems. Based on this analysis, we determined that DMA and rheology provide the broadest capability while remaining robust and easy to use. Other techniques, such as nanoindentation and vibrations, have unique qualities that fulfill niche applications where DMA and rheology are not suitable. This analysis provides an industry-relevant evaluation of measurement techniques and demonstrates a framework for evaluating the capabilities of analytical equipment relative to organizational needs.