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Convergent beam electron diffraction extinction distance measurements for quantitative analysis of si(1-x),Ge(x).

A new method to determine the concentration of germanium in Si(1-x) Ge(x) single crystals is presented. It is based on extinction distance measurements by means of convergent beam electron diffraction (CBED). The two-beam condition CBED intensity oscillation (the so-called rocking curve) is measured for the 004 diffracted beam and compared with a numerical simulation. Using the two-beam dynamical diffraction approximation theory, this approach yields very precise values for both specimen thickness and effective extinction distance (Ultramicroscopy 87 (2001) 5). First a theoretical extinction distance zetag(x) for strain relaxed Si(1-x)Ge(x) is calculated assuming a solid solution and using tabulated atomic scattering factors of silicon and germanium atoms. It is found that for single crystals zetag(x) decreases from 156 nm in pure silicon to 90 nm in pure germanium. Measurements on calibrated strain relaxed SiGe layers with variable germanium concentrations show an excellent agreement between experimental and calculated extinction distances zetag(x). As a consequence the experimental extinction distance zetag(x) becomes an indirect measure of the germanium concentration with a 1-2 at % sensitivity. The method turns out to be insensitive to strain as experimental zetag(x) values obtained on strained SiGe layers fit the theoretical extinction distance curve calculated for strain relaxed SiGe.

Journal Article↗

Cytotoxic lesion of the medial prefrontal cortex abolishes the partial reinforcement extinction effect, attenuates prepulse inhibition of the acoustic startle reflex and induces transient hyperlocomotion, while sparing spontaneous object recognition memory in the rat.

The partial reinforcement extinction effect refers to the increase in resistance to extinction of an operant response acquired under partial reinforcement relative to that acquired under continuous reinforcement. Prepulse inhibition of the acoustic startle response refers to the reduction in startle reactivity towards an intense acoustic pulse stimulus when it is shortly preceded by a weak prepulse stimulus. These two behavioural phenomena appear to be related to different forms of attentional processes. While the prepulse inhibition effect reflects an inherent early attentional gating mechanism, the partial reinforcement extinction effect is believed to involve the development of acquired inattention, i.e. the latter requires the animals to learn about what to and what not to attend. Impairments in prepulse inhibition and the partial reinforcement extinction effect have been independently linked to the neuropsychology of attentional dysfunctions seen in schizophrenia. The proposed neural substrates underlying these behaviourial phenomena also appear to overlap considerably: both focus on the nucleus accumbens and emphasize the functional importance of its limbic afferents, including that originating from the medial prefrontal cortex, on accumbal output/activity. The present study demonstrated that cytotoxic medial prefrontal cortex lesions which typically damaged the prelimbic, the infralimbic and the dorsal anterior cingulate areas could lead to the abolition of the partial reinforcement extinction effect and the attenuation of prepulse inhibition. The lesions also resulted in a transient elevation of spontaneous locomotor activity. In contrast, the same lesions spared performance in a spontaneous object recognition memory test, in which the lesioned animals displayed normal preference for a novel object when the novel object was presented in conjunction with a familiar object seen 10 min earlier within an open field arena. The present results lend support to the hypothesis that medial prefrontal cortex dysfunction might be related to some forms of attentional abnormality central to the symptomatology of schizophrenia. Relevance of the present findings in relation to the neural substrates underlying the partial reinforcement extinction effect and prepulse inhibition is further discussed.

Acoustic Stimulation↗

Superstitious responding and reinforcement rate under concurrent variable-interval extinction schedules.

To examine superstitious responding, four pigeons key pecked under multiple concurrent variable-interval 45 s variable-interval 90 s concurrent variable-interval 90 s variable-interval 180 s schedules in the absence of a changeover delay. The two variable-interval 90 s schedules then were replaced by extinction, and key-peck responding during extinction was examined as a function of the prevailing reinforcement rate. During the first several sessions, extinction-key responding was maintained closer to baseline levels in the presence of the higher reinforcement rate, and this effect dissipated or even reversed with continued exposure to extinction. Although extinction-key responding generally decreased to near-zero levels after several sessions, in a few instances, it continued for 30 and 45 sessions. These results demonstrate how concurrent variable-interval extinction schedules can be used to investigate what often has been labeled superstitious responding.

Journal Article↗

Extinction from a paleontological perspective.

Extinction of widespread species is common in evolutionary time (millions of years) but rare in ecological time (hundreds or thousands of years). In the fossil record, there appears to be a smooth continuum between background and mass extinction; and the clustering of extinctions at mass extinctions cannot be explained by the chance coincidence of independent events. Although some extinction is selective, much is apparently random in that survivors have no recognizable superiority over victims. Extinction certainly plays an important role in evolution, but whether it is constructive or destructive has not yet been determined.

Adaptation, Biological↗

Rapid recovery from the Late Ordovician mass extinction.

Understanding the evolutionary role of mass extinctions requires detailed knowledge of postextinction recoveries. However, most models of recovery hinge on a direct reading of the fossil record, and several recent studies have suggested that the fossil record is especially incomplete for recovery intervals immediately after mass extinctions. Here, we analyze a database of genus occurrences for the paleocontinent of Laurentia to determine the effects of regional processes on recovery and the effects of variations in preservation and sampling intensity on perceived diversity trends and taxonomic rates during the Late Ordovician mass extinction and Early Silurian recovery. After accounting for variation in sampling intensity, we find that marine benthic diversity in Laurentia recovered to preextinction levels within 5 million years, which is nearly 15 million years sooner than suggested by global compilations. The rapid turnover in Laurentia suggests that processes such as immigration may have been particularly important in the recovery of regional ecosystems from environmental perturbations. However, additional regional studies and a global analysis of the Late Ordovician mass extinction that accounts for variations in sampling intensity are necessary to confirm this pattern. Because the record of Phanerozoic mass extinctions and postextinction recoveries may be compromised by variations in preservation and sampling intensity, all should be reevaluated with sampling-standardized analyses if the evolutionary role of mass extinctions is to be fully understood.

Biodiversity↗

Lessons from the past: evolutionary impacts of mass extinctions.

Mass extinctions have played many evolutionary roles, involving differential survivorship or selectivity of taxa and traits, the disruption or preservation of evolutionary trends and ecosystem organization, and the promotion of taxonomic and morphological diversifications-often along unexpected trajectories-after the destruction or marginalization of once-dominant clades. The fossil record suggests that survivorship during mass extinctions is not strictly random, but it often fails to coincide with factors promoting survival during times of low extinction intensity. Although of very serious concern, present-day extinctions have not yet achieved the intensities seen in the Big Five mass extinctions of the geologic past, which each removed > or =50% of the subset of relatively abundant marine invertebrate genera. The best comparisons for predictive purposes therefore will involve factors such as differential extinction intensities among regions, clades, and functional groups, rules governing postextinction biotic interchanges and evolutionary dynamics, and analyses of the factors that cause taxa and evolutionary trends to continue unabated, to suffer setbacks but resume along the same trajectory, to survive only to fall into a marginal role or disappear ("dead clade walking"), or to undergo a burst of diversification. These issues need to be addressed in a spatially explicit framework, because the fossil record suggests regional differences in postextinction diversification dynamics and biotic interchanges. Postextinction diversifications lag far behind the initial taxonomic and morphological impoverishment and homogenization; they do not simply reoccupy vacated adaptive peaks, but explore opportunities as opened and constrained by intrinsic biotic factors and the ecological and evolutionary context of the radiation.

Animals↗

Genetic variability and effective population size when local extinction and recolonization of subpopulations are frequent.

If a population (species) consists of n haploid lines (subpopulations) which reproduce asexually and each of which is subject to random extinction and subsequent replacement, it is shown that, at equilibrium in which mutational production of new alleles and their random extinction balance each other, the genetic diversity (1 minus the sum of squares of allelic frequencies) is given by 2N(e)v/(1 + 2N(e)v), where [Formula: see text] in which N is the harmonic mean of the population size per line, n is the number of lines (assumed to be large), lambda is the rate of line extinction, and v is the mutation rate (assuming the infinite neutral allele model). In a diploid population (species) consisting of n colonies, if migration takes place between colonies at the rate m (the island model) in addition to extinction and recolonization of colonies, it is shown that effective population size is [Formula: see text] If the rate of colony extinction (lambda) is much larger than the migration rate of individuals, the effective population size is greatly reduced compared with the case in which no colony extinctions occur (in which case N(e) = nN). The stepping-stone type of recolonization scheme is also considered. Bearing of these results on the interpretation of the level of genetic variability at the enzyme level observed in natural populations is discussed from the standpoint of the neutral mutation-random drift hypothesis.

Journal Article↗

Radiocarbon dates on bones of extinct birds from Hawaii.

Bones from a stratified sedimentary deposit in the Puu Naio Cave site on Maui, Hawaiian Islands, reveal the late Holocene extinction of 19 species of birds. The age of the sediment and associated fauna was determined by direct radiocarbon dating (tandem particle accelerator-mass spectrometer; TAMS) of amino acids extracted from bones weighing as little as 450 mg. The 14C dates indicate that sediment has been accumulating in the lava tube for at least the last 7750 years, a suitable time frame for testing the hypothesis that Holocene extinction on islands began after human colonization. Despite growing evidence that a worldwide wave of extinctions coincided with human colonization of oceanic islands, little radiometric data have been available to date the extinction of most small fossil vertebrates on islands. The TAMS technique of dating purified collagen from the bones of small vertebrates could lead to vastly improved chronologies of extinction for oceanic islands where catastrophic mid- to late-Holocene extinction is expected or known to have occurred. Chronologies derived from nonarcheological sites that show continuous sedimentation, such as the Puu Naio Cave deposit, may also yield key evidence on the timing of earliest human settlement of Oceania.

Animals↗

Extinction of JC virus tumor-antigen expression in glial cell--fibroblast hybrids.

JC virus (JCV) is a ubiquitous human papovavirus that shares sequence and structural homology with simian virus 40 (SV40). In contrast to SV40, expression of JCV is restricted to a small number of cell types, including human fetal glial cells, uroepithelial cells, amnion cells, and some endothelial cells. To study the control of JCV early region expression, we made heterokaryons and stable hybrids between JCV-transformed hamster glial cells and mouse fibroblasts. Binucleate heterokaryons exhibited extinction of large tumor antigen expression in the hamster nuclei as assayed by indirect immunofluorescence. This extinction was both time and dose dependent: extinction reached maximal levels at 24-36 hr after fusion and was dependent on the ratio of glial cell to fibroblast nuclei in multinucleated heterokaryons. Extinction also was observed in stable hybrids between the glial cells and mouse Ltk- cells. Southern blot analysis showed that the extinguished hybrids contained viral sequences. Reexpression of large tumor antigen was observed in several subclones, suggesting that extinction was correlated with the loss of murine fibroblast chromosomes from these hybrids. The cis-acting region that mediates extinction resides within the viral regulatory region, which contains two 98-base-pair repeats that have enhancer activity. These data demonstrate that cellular factors that negatively regulate viral gene expression contribute to the restricted cell-type specificity of this virus.

Animals↗

Unusual dynamics of extinction in a simple ecological model.

Studies on natural populations and harvesting biological resources have led to the view, commonly held, that (i) populations exhibiting chaotic oscillations run a high risk of extinction; and (ii) a decrease in emigration/exploitation may reduce the risk of extinction. Here we describe a simple ecological model with emigration/depletion that shows behavior in contrast to this. This model displays unusual dynamics of extinction and survival, where populations growing beyond a critical rate can persist within a band of high depletion rates, whereas extinction occurs for lower depletion rates. Though prior to extinction at lower depletion rates the population exhibits chaotic dynamics with large amplitudes of variation and very low minima, at higher depletion rates the population persists at chaos but with reduced variation and increased minima. For still higher values, within the band of persistence, the dynamics show period reversal leading to stability. These results illustrate that chaos does not necessarily lead to population extinction. In addition, the persistence of populations at high depletion rates has important implications in the considerations of strategies for the management of biological resources.

Animals↗

Late quaternary extinction of a tree species in eastern North America.

Widespread species- and genus-level extinctions of mammals in North America and Europe occurred during the last deglaciation [16,000-9,000 yr B.P. (by (14)C)], a period of rapid and often abrupt climatic and vegetational change. These extinctions are variously ascribed to environmental change and overkill by human hunters. By contrast, plant extinctions since the Middle Pleistocene are undocumented, suggesting that plant species have been able to respond to environmental changes of the past several glacial/interglacial cycles by migration. We provide evidence from morphological studies of fossil cones and anatomical studies of fossil needles that a now-extinct species of spruce (Picea critchfieldii sp. nov.) was widespread in eastern North America during the Last Glacial Maximum. P. critchfieldii was dominant in vegetation of the Lower Mississippi Valley, and extended at least as far east as western Georgia. P. critchfieldii disappeared during the last deglaciation, and its extinction is not directly attributable to human activities. Similarly widespread plant species may be at risk of extinction in the face of future climate change.

Biological Evolution↗

Life-history models of extinction: a test with island spiders.

This study analyzes extinction patterns for two species of orb spiders monitored annually on 77 islands over a continuous 20-yr period. One species, Argiope argentata, has large populations sometimes crashing quickly to extinction and a much weaker relation of extinction likelihood to population size than does the other species, Metepeira datona. Demographic models were built for both species and matched against observations. Differences between the species in life-history traits-estimated with measurements from the field-together with incorporation of demographic stochasticity, a population ceiling, and environmental stochasticity, were necessary to fit the observed extinction curves. As predicted from life-history patterns, long-term population growth rates (and hence predicted extinction probabilities) are relatively very sensitive to values of juvenile survivorship. Models are also sensitive to variation in the population ceiling and environmental noise, which tend to act in a complementary manner. A simple model with no age structure was able to fit the data on large initial population sizes but not on small initial population sizes, showing that life cycle characteristics interact with the various sources of stochasticity and hence have to be taken into account to produce a precise model of the extinction process.

Age Factors↗

Selective extinction and rapid loss of evolutionary history in the bird fauna.

The extinction of species results in a permanent loss of evolutionary history. Recent theoretical studies show that this loss may be proportionally much smaller than the loss of species, but under some conditions can exceed it. Such conditions occur when the phylogenetic tree that describes the evolutionary relationships among species is highly imbalanced due to differences between lineages in past speciation and/or extinction rates. I used the taxonomy by C. G. Sibley and B. L. Monroe Jr to estimate the global loss of bird evolutionary history from historical and predicted extinctions, and to quantify the ensuing changes in balance of the bird phylogenetic tree. In the global bird fauna, evolutionary history is being lost at a high rate, similar to the rate of species extinction. The bird phylogenetic tree is highly imbalanced, and the imbalance is increased significantly by anthropogenic extinction. Historically, the elevated loss of bird evolutionary history has been fuelled mostly by phylogenetic non-randomness in the extinction of species, but the direct effect of tree imbalance is substantial and could dominate in the future.

Animals↗

Human demography and reserve size predict wildlife extinction in West Africa.

Species-area models have become the primary tool used to predict baseline extinction rates for species in isolated habitats, and have influenced conservation and land-use planning worldwide. In particular, these models have been used to predict extinction rates following the loss or fragmentation of natural habitats in the absence of direct human influence on species persistence. Thus, where direct human influences, such as hunting, put added pressure on species in remnant habitat patches, we should expect to observe extinction rates higher than those predicted by simple species-area models. Here, we show that extinction rates for 41 species of large mammals in six nature reserves in West Africa are 14-307 times higher than those predicted by models based on reserve size alone. Human population and reserve size accounted for 98% of the observed variation in extinction rates between reserves. Extinction occurred at higher rates than predicted by species-area models for carnivores, primates and ungulates, and at the highest rates overall near reserve borders. Our results indicate that, where the harvest of wildlife is common, conservation plans should focus on increasing the size of reserves and reducing the rate of hunting.

Animals↗

Sexual selection and the risk of extinction in birds.

The relationship between sexual selection and extinction risk has rarely been investigated. This is unfortunate because extinction plays a key role in determining the patterns of species richness seen in extant clades, which form the basis of comparative studies into the role that sexual selection may play in promoting speciation. We investigate the extent to which the perceived risk of extinction relates to four different estimates of sexual selection in 1030 species of birds. We find no evidence that the number of threatened species is distributed unevenly according to a social mating system, and neither of our two measures of pre-mating sexual selection (sexual dimorphism and dichromatism) was related to extinction risk, after controlling for phylogenetic inertia. However, threatened species apparently experience more intense post-mating sexual selection, measured as testis size, than non-threatened species. These results persisted after including body size as a covariate in the analysis, and became even stronger after controlling for clutch size (two known correlates of extinction risk). Sexual selection may therefore be a double-edged process-promoting speciation on one hand but promoting extinction on the other. Furthermore, we suggest that it is post-mating sexual selection, in particular, that is responsible for the negative effect of sexual selection on clade size. Why this might be is unclear, but the mean population fitness of species with high intensities of post-mating sexual selection may be especially low if costs associated with multiple mating are high or if the selection load imposed by post-mating selection is higher relative to that of pre-mating sexual selection.

Animals↗

Evolution and extinction in the marine realm: some constraints imposed by phytoplankton.

The organic and mineralized remains of planktonic algae provide a rich record of microplankton evolution extending over nearly half of the preserved geological record. In general, Phanerozoic patterns of phytoplankton radiation and extinction parallel those documented for skeletonized marine invertebrates, both augmenting and constraining thought about evolution in the oceans. Rapidly increasing knowledge of Proterozoic plankton is making possible the recognition of additional episodes of diversification and extinction that antedate the Ediacaran radiation of macroscopic animals. In contrast to earlier phytoplankton history, the late Mesozoic and Cainozoic record is documented in sufficient detail to constrain theories of mass extinction in more than a general way. Broad patterns of diversity change in planktonic algae show similarities across the Cretaceous-Tertiary and Eocene-Oligocene boundaries, but detailed comparisons of origination and extinction rates in calcareous nannoplankton, as well as other algae and skeletonized protozoans, suggest that the two episodes were quite distinct. Common causation appears unlikely, casting doubt on monolithic theories of mass extinction, whether periodic or not. Studies of mass extinction highlight a broader class of insights that paleontologists can contribute to evolutionary biology: the evaluation of evolutionary change in the context of evolving Earth-surface environments.

Biological Evolution↗

Diversification and extinction patterns among Neogene perimediterranean mammals.

The best mammalian fossil record during the Neogene of Western Europe is that of the rodents, the most successful and diversified mammal order. The study of origination and extinction during the Neogene (24-3 Ma BP) in one of the best-documented areas, Spain and southern France, gives an insight into the dynamics of these communities and indicates the possible nature of the driving forces. Three main periods of time show a high rate of origination: the late Burdigalian (17.5 Ma BP), the early Vallesian (11.5-11 Ma BP) and the early Pliocene (4.2-3.8 Ma BP). Two of these high origination-rate periods are immediately followed by important extinction events during which all cohorts are deeply affected (11.5-11 Ma BP and 4.2-3.8 Ma BP). The most important extinction event seems to occur during the early Vallesian (11.5-11 Ma BP), which probably includes the middle/late Miocene boundary. At the Miocene/Pliocene boundary, and during the early Pliocene, the faunal turnover seems to become faster, inducing a strong decrease of the mean species duration. Whereas the main immigration event, which occurs at 17.5 Ma BP, can be related to other faunal migrations in terms of the closure of the Tethys, as it occurs also in eastern Africa and in southwest Asia, the middle/late Miocene boundary event may have been related to a period of ice growth in the Southern Hemisphere. The extinction event that affects the planktonic foraminifera at 12 Ma BP cannot be chronologically correlated to this southwestern European land-mammal extinction event, because the calibration of the marine fossil record during that time-span has to be precise. Some limited terrestrial faunal exchanges that occur during the Messinian between southwestern Europe and northwestern Africa do not deeply affect the general faunal dynamics. Both allochthonous cohorts of immigrants become rapidly extinct. Several endemic rodent faunas, indicating insular conditions, have been reported from the southern edge of the western European continent from the middle Miocene up to the Pliocene. All show low taxonomic diversity, strong endemism and short survival. Some of them, like those of the Gargano Islands during the late Miocene, underwent peculiar morphological changes and also speciation. The large number of rodent genera coevolving in the Gargano Islands is indicative of the large surface areas of these islands. The general geographic pattern of southwestern Europe during the Neogene may therefore correspond to a large continental province including Spain and southern France with some kind of fast-modifying archipelago on its southern rim.

Animals↗

Natural extinction on islands.

Almost all recent extinction of species or subspecies on islands comes from human activities. On the other hand, in local populations there is much natural extinction and immigration, i.e. turnover, on small islands. Most of this turnover occurs in locally rare species, and attests to the phenomenon of minimum viable population size. The MacArthur-Wilson theory is based on this turnover which, from an ecological point of view, is generally trivial. More useful theories of minimum viable population size are being developed. Rarity is the precursor of extinction, and species can be rare in several ways. Models of these phenomena are still primitive, particularly those that relate habitat availability to population density. Models of interactive communities show phenomena that may be relevant to the understanding of extinction in the geological record. Lotka-Volterra equations indicate considerable sensitivity to invasions, sometimes producing a cascade of extinction. Chemostat equations show that the behaviour of food chains can change dramatically with small changes in parameters, suggesting that small environmental effects can sometimes cause large ecological changes, including extinctions, in interactive biotic communities.

Animals↗