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Diversity components of impending primate extinctions.

Many extant species are at risk to go extinct. This impending loss of species is likely to cause changes in future ecosystem functions. Ecological components of diversity, such as dietary or habitat specializations, can be used to estimate the impact of extinctions on ecosystem functions. As an approach to estimate the impact of future extinctions, we tested interdependency between ecological and taxonomic change based on current predictions of extinction rates in primates. We analyzed the ecological characteristics of extant primate faunas having species in various categories of endangerment of extinction and forecasted the future primate faunas as if they were paleontological faunas. Predicting future faunas combines the wealth of ecological information on living primates with large, fossil record-like changes in diversity. Predicted extinction patterns of living primates in Africa, Asia, Madagascar, and South America show that changes in ecology differ among the regions in ways that are not reducible to taxonomic measures. The ecological effects of primate extinctions are initially least severe in South America and larger in Asia and Africa. Disproportionately larger ecological changes are projected for Madagascar. The use of taxonomy as a proxy for ecology can mislead when estimating competence of future primate ecosystems.

Animals↗

Back-casting sociality in extinct species: new perspectives using mass death assemblages and sex ratios.

Despite 150 years of interest in the ecology of dinosaurs, mammoths, proto-hominids and other extinct vertebrates, a general framework to recreate patterns of sociality has been elusive. Based on our recent discovery of a contemporary heterospecific mass death assemblage in the Gobi Desert (Mongolia), we fit predictions about gender-specific associations and group living in extant ungulates to extinct ones. We relied on comparative data on sex-ratio variation and body-size dimorphism, basing analyses on 38 additional mass mortality sites from Asia, Africa, Europe and North America that span 50 million years. Both extant and extinct species died in aggregations with biased adult sex ratios, but the skew (from 1:1) was greater for extinct dimorphic taxa, suggesting that sociality in these extinct species can be predicted from spatial and demographic traits of extant ones. However, extinct rhinos, horses and zebras were inconsistent with predictions about adult sex ratios, which underscores the inherent difficulty in backcasting historic patterns to some monomorphic taxa. These findings shed light not only on the sociality of extinct species but provide a sound, although limited, footing for interpretation of modern death assemblages within the context of the emerging science of taphonomy and palaeobehaviour.

Animals↗

Character release following extinction in a Caribbean reef coral species complex.

The Pleistocene extinction of the widespread organ-pipe Montastraea coral had measurable morphological and ecological effects on surviving lineages of the Montastraea "annularis" species complex. Extinction of the organ-pipe Montastraea occurred after more than 500,000 years of dominance in the shallow-water reef habitat of Barbados. Extinction resulted in a morphological shift of the columnar Montastraea lineage from thick to thin columns in modern reef environments. Pleistocene colonies of the columnar morphotype sympatric with organ-pipe Montastraea showed greater column widths than those in allopatry. We subjected our data to a number of criteria for interpreting the morphological shift as character release following lifting of competitive pressure after extinction. The morphological differences do not appear to be due either to chance or to physical properties of the marine environment. Differential local extinction and recolonization of four members of the species complex did not occur on Barbados, so that the species coexisted and appear to have coevolved between more than 600,000 and 82,000 years ago. The morphological shift is related to coral growth form and growth rate, and thus reflects the acquisition of a primary resource in corals--light. Character release occurred at the same oceanic Caribbean island (Barbados) where environments have fluctuated with similar variance throughout the period of coexistence. Not only has competition among living members of the Montastraea "annularis" species complex been convincingly demonstrated, but trends in relative abundance among fossil members of the species complex strongly suggest that a competitive hierarchy was operating during their Pleistocene coexistence on Barbados. We also observed an ecological analogue to character release on another Caribbean island. Curaçao. The distribution and abundance of living columnar M. annularis s.s. and massive M. faveolata from the leeward reef crest in Curaçao is greater now than in the Pleistocene, when organ-pipe Montastraea dominated this shallow-water reef habitat. Extinction of the faster growing, shallow-water organ-pipe Montastraea resulted in higher abundance of the columnar Montastraea lineage in shallow-water habitats, where it shifted its morphology to one adapted to high light levels. The species extinction released surviving lineages from a competitive network that had resulted in lower rank abundance in the Pleistocene community and enhanced abundance of both columnar M. annularis s.s. and M. faveolata in modern communities. Full validation of our interpretation of character release must await experiments that demonstrate whether phenotypic differences between populations have a genetic basis. However, we believe the results of this study point to the important, yet heretofore neglected, role that biological interactions have played in the evolution of closely related reef coral species.

Animals↗

Pharmacological manipulations of the extinction process of fear-induced ultrasonic vocalization in rats.

The effects of pharmacological manipulation on the extinction process of fear-induced ultrasonic vocalizations (USVs), which are considered distress calls related to anxiety, were investigated. Male Wistar rats were conditioned to emit USVs by being given repeated electrical foot-shocks while in a chamber. After 10 sessions of conditioning, the animals started to emit USVs upon mere exposure to the shock chamber without being shocked. Using these animals, the extinction process of the USVs was examined. With repeated exposure to the chamber without shocks, the USVs first increased and then gradually decreased. i.e., the extinction burst was observed. Daily intraperitoneal injections of a benzodiazepine-GABA receptor agonist diazepam (DZP; 1.0 mg/kg) or a tricyclic antidepressant clomipramine (CLM; 20 mg/kg) inhibited this extinction burst. Moreover, CLM, but not DZP, shortened the period required for extinction as compared with the vehicle-treated animals. Following the extinction phase, the emission of USVs was enhanced by the cessation of both drug treatments. These results suggest that CLM would be useful for reducing anxiety-related behaviors in the extinction process, as long as withdrawal symptoms after long-term drug treatments are taken into consideration.

Animals↗

Human population density and extinction risk in the world's carnivores.

Understanding why some species are at high risk of extinction, while others remain relatively safe, is central to the development of a predictive conservation science. Recent studies have shown that a species' extinction risk may be determined by two types of factors: intrinsic biological traits and exposure to external anthropogenic threats. However, little is known about the relative and interacting effects of intrinsic and external variables on extinction risk. Using phylogenetic comparative methods, we show that extinction risk in the mammal order Carnivora is predicted more strongly by biology than exposure to high-density human populations. However, biology interacts with human population density to determine extinction risk: biological traits explain 80% of variation in risk for carnivore species with high levels of exposure to human populations, compared to 45% for carnivores generally. The results suggest that biology will become a more critical determinant of risk as human populations expand. We demonstrate how a model predicting extinction risk from biology can be combined with projected human population density to identify species likely to move most rapidly towards extinction by the year 2030. African viverrid species are particularly likely to become threatened, even though most are currently considered relatively safe. We suggest that a preemptive approach to species conservation is needed to identify and protect species that may not be threatened at present but may become so in the near future.

Animals↗

Extinction debt of forest plants persists for more than a century following habitat fragmentation.

Following habitat fragmentation individual habitat patches may lose species over time as they pay off their "extinction debt." Species with relatively low rates of population extinction and colonization ("slow" species) may maintain extinction debts for particularly prolonged periods, but few data are available to test this prediction. We analyzed two unusually detailed data sets on forest plant distributions and land-use history from Lincolnshire, United Kingdom, and Vlaams-Brabant, Belgium, to test for an extinction debt in relation to species-specific extinction and colonization rates. Logistic regression models predicting the presence-absence of 36 plant species were first parameterized using data from Lincolnshire, where forest cover has been relatively low (approximately 5-8%) for the past 1000 years. Consistent with extinction debt theory, for relatively slow species (but not fast species) these models systematically underpredicted levels of patch occupancy in Vlaams-Brabant, where forest cover was reduced from approximately 25% to <10% between 1775 and 1900 (it is presently 6.5%). As a consequence, the ability of the Lincolnshire models to predict patch occupancy in Vlaams-Brabant was worse for slow than for fast species. Thus, more than a century after forest fragmentation reached its current level an extinction debt persists for species with low rates of population turnover.

Belgium↗

Effects of methylphenidate on responding under extinction in the presence and absence of conditioned reinforcement.

Pigeons pecked a key during sessions that began with a variable number of reinforcers under a second-order schedule of food presentation. Every 30sec, on the average, a key peck was followed immediately by one of two consequences: (a) food presentation, accompanied by a stimulus complex that consisted of houselight off, key color change, tone presentation, and hopper-light illumination, or (b) the stimulus complex alone. Following the last food presentation, 20min of one of two types of extinction began. The two types of extinction were: (a) standard extinction (key pecks had no consequence) and (b) key pecks produced, on a variable-interval schedule, the stimulus complex previously paired with food. Consequently, it was possible to study performance under extinction during which responses either did or did not result in occasional presentation of a food-paired stimulus complex. Methylphenidate (5, 10 and 20mg/kg) occasionally was administered before sessions containing each type of extinction. At moderate doses methylphenidate produced higher response rates during extinction when the stimulus complex was presented than when it was not. These results support previous findings with rats that stimulant drugs can enhance responding during extinction when responding produces conditioned reinforcers, and illustrate this effect in a novel, within-subject design.

Journal Article↗

The case for extraterrestrial causes of extinction.

The dramatic increase in our knowledge of large-body impacts that have occurred in Earth's history has led to strong arguments for the plausibility of meteorite impact as a cause of extinction. Proof of causation is often hampered, however, by our inability to demonstrate the synchronism of specific impacts and extinctions. A central problem is range truncation: the last reported occurrences of fossil taxa generally underestimate the true times of extinction. Range truncation, because of gaps in sedimentation, lack of preservation, or lack of discovery, can make sudden extinctions appear gradual and gradual extinctions appear sudden. Also, stepwise extinction may appear as an artefact of range truncation. These effects are demonstrated by experiments performed on data from field collections of Cretaceous ammonities from Zumaya (Spain). The challenge for future research is to develop a new calculus for treating biostratigraphic data so that fossils can provide more accurate assessments of the timing of extinctions.

Animals↗

The case for extraterrestrial causes of extinction.

The dramatic increase in our knowledge of large-body impacts that have occurred in Earth's history has led to strong arguments for the plausibility of meteorite impact as a cause of extinction. Proof of causation is often hampered, however, by our inability to demonstrate the synchronism of specific impacts and extinctions. A central problem is range truncation: the last reported occurrences of fossil taxa generally underestimate the true times of extinction. Range truncation, because of gaps in sedimentation, lack of preservation, or lack of discovery, can make sudden extinctions appear gradual and gradual extinctions appear sudden. Also, stepwise extinction may appear as an artefact of range truncation. These effects are demonstrated by experiments performed on data from field collections of Cretaceous ammonities from Zumaya (Spain). The challenge for future research is to develop a new calculus for treating biostratigraphic data so that fossils can provide more accurate assessments of the timing of extinctions.

Animals↗

Context extinction and associative learning in Lymnaea.

Aerial respiratory behavior in the pond snail Lymnaea was operantly conditioned so that snails learned not to perform aerial respiration in a hypoxic environment. Snails were trained in either the standard context (no food odorant) or a carrot (food-odorant) context. An operant training procedure of two 45-min training sessions with a 1-h interval between the sessions followed by a third 45-min training session 18 h later was sufficient to produce associative learning and long-term memory (LTM) that persisted for at least 5 days. If, however, following the third operant training session snails received three 45-min extinction training sessions, with each extinction session separated by at least a 1-h interval, LTM was not observed when tested the following day. That is, the memory was extinguished. Extinction, however, did not occur if the context of the extinction training was different from the context of the associative training. That is, in the snails trained in the standard context, extinction did not occur if the extinction training sessions were performed in the food-odorant context and vice versa.

Animals↗

The effect of morphine on fear extinction in rats.

Rats were trained on an appetitive discretetrial discriminated-punishment task in which they learned to suppress responding when an intense flashing light predicting punishment was present and to respond rapidly on trials when the flashing light was absent. Once animals were performing discriminatively, 0.75, 3.0, or 6.0 mg/kg of morphine (base) was administered and a fear extinction session consisting of 60 nonshocked presentations of the flashing light was given. Two saline control groups, one that received fear extinction and one that did not, were also included in the experiment. On the day following fear extinction, all rats were tested in the undrugged state on the discriminated punishment problem, but without shock. The rats receiving 3.0 and 6.0 mg/kg of morphine before the fear extinction session were suppressed by the flashing light more than the saline extinction group or the 0.75 mg/kg morphine treatment group. Moreover, the two higher dose morphine groups were suppressed as readily as the saline group that received no fear extinction. These results are attributed to the antiemotionality effects of morphine.

Animals↗

Extinction and dopamine receptor blockade after intermittent reinforcement training: failure to observe functional equivalence.

Response decrements in an operant task produced by either extinction or by the dopamine receptor blocker pimozide were examined in three experiments which employed intermittent reinforcement schedules. In contrast to the congruency between these treatments previously observed following continuous reinforcement training, treatment with pimozide was markedly more effective than extinction in decreasing performance after training with variable interval, fixed interval, and fixed ratio reinforcement. The two treatments also produced substantially different patterns of responding. A shift from extinction to pimozide did not alter the progressive decline in response rate over days, but a shift from pimozide to extinction caused a pronounced increase of performance. These results indicate that the pimozide and extinction treatment did not produce functionally equivalent effects, and that the role of dopamine on reward processes should not be inferred from comparisons between pimozide and extinction.

Animals↗

Reduced behavioral variability in extinction: effects of chronic treatment with the benzodiazepine, diazepam or with ethanol.

Extinction of a food reinforced habit results in an increase in the variability of the response learned in acquisition and in the appearance of previously suppressed competing responses. The purpose of the present study was to examine the effects of chronically administered diazepam (0.0, 1.5, 3.0, or 6.0 mg/kg, IP, -30 min) or 10% ethanol (0.0, 1.0, 1.5, or 2.0 g/kg, IP, -15 min) on such behavioral variability in the extinction of radial maze performance. Eight groups of food deprived rats (n = 6) were given one of the forementioned doses for 2 sessions of baseline, 18 sessions of acquisition, and 5 sessions of extinction. In acquisition, eight rewards of two food pellets were obtained on each of three trials in each session. The food well at the end of each arm was rebaited when emptied by the animal, consequently an entry into any arm was reinforced. In baseline and extinction no food was available in the maze. Each session consisted of three 10-min trials. In extinction, compared to treatment with vehicle, both diazepam and ethanol treatments decreased the rate of the instrumental response, arm entry, and increased the variability of the instrumental response and of competing responses. Only the effects of the drugs on the competing responses in extinction were greater than those observed in acquisition. It was concluded that the interference-reduction model of drug action best described the magnitude of the drug effects and the variability-reduction model best predicted the direction of the effects.

Animals↗

Revisiting the learning-without-awareness question in human Pavlovian autonomic conditioning: focus on extinction in a dichotic listening paradigm.

Numerous studies have indicated that, consistent with current "cognitive" accounts of information processing, human Pavlovian autonomic discrimination acquisition cannot occur without awareness of the CS-US relationship. However, extinction studies have suggested that awareness is not necessary, findings that, in information-processing terms, have been explained by assuming that the processing by the extinction stage is parallel (automatic) rather than serial (controlled). This explanation was tested in an 80-subject study. The first, acquisition phase was a standard semantic differential conditioning arrangement with a 96-db white noise as US, and a "long" CS-US interval of 8 s, with ten trials each of CS+ (paired with US) and CS- (unpaired) trials. In extinction (USs omitted), in order to obtain non-autonomic indices of processing and thereby test the information-processing account of "unaware" autonomic conditioning during extinction, a dichotic listening task was implemented, with the CSs presented in the unattended channel (ear), while the subject had to perform a semantic differential reaction task in an attended-to channel (other ear). In early extinction, the electrodermal response occurring at an interval of 9-15 s after CS onset (i.e., following placement of the US during acquisition) and the finger-pulse-volume response occurring at an interval of 4-11 s after CS onset both showed reliable conditioning, but reaction-time and subjective-report data for the recognized critical words indicated serial rather than parallel processing of the CSs during extinction.

Adult↗

Effect of low doses of delta9-tetrahydrocannabinol and cannabidiol on the extinction of cocaine-induced and amphetamine-induced conditioned place preference learning in rats.

RATIONALE: Using the place-preference conditioning paradigm, we evaluated the potential of the two most prominent cannabinoids found in marijuana, the psychoactive component delta9-tetrahydrocannabinol (delta9-THC) and the nonpsychoactive component cannabidiol (CBD), to potentiate extinction of a cocaine-induced and an amphetamine-induced conditioned place preference in rats. METHODS: To determine the effects of pretreatment with delta9-THC or CBD on extinction, a cocaine-induced and amphetamine-induced place preference was first established. Rats were then given an extinction trial, during which they were confined to the treatment-paired floor for 15 min. Thirty minutes prior to the extinction trial, they were injected with a low dose of delta9-THC (0.5 mg/kg), CBD (5 mg/kg) or vehicle. The potential of the CB1 receptor antagonist, SR141716, to reverse the effects of delta9-THC or CBD was also evaluated. To determine the hedonic effects of CBD, one distinctive floor was paired with CBD (5 mg/kg) and another with saline. Finally, to determine the effect of delta9-THC.or CBD on the establishment and/or the expression of a place preference during four cycles of conditioning trials, rats were injected with delta9-THC (0.25-1 mg/kg), CBD (5 mg/kg) or vehicle 25 min prior to receiving an injection of amphetamine followed by placement on the treatment floor; on alternate days, they received injections of vehicle followed by saline and placement on the nontreatment floor. The rats then received two test trials; on one trial they were pretreated with the cannabinoid and on the other trial with vehicle. RESULTS: delta9-THC and CBD potentiated the extinction of both cocaine-induced and amphetamine-induced conditioned place preference learning, and this effect was not reversed by SR141716. The cannabinoids did not affect learning or retrieval, and CBD was not hedonic on its own. CONCLUSIONS: These results are the first to show that both delta9-THC, which acts on both CB 1 and CB2 receptors, and CBD, which does not bind to CB1 or CB2 receptors, potentiate the extinction of conditioned incentive learning.

Amphetamine↗

Evidence that GABA transmission mediates context-specific extinction of learned fear.

Six experiments used rats to study the effects of the beta-carboline FG 7142 on extinction of fear responses (freezing) to an auditory cue that had signalled footshock. Subcutaneous injection of FG 7142 interfered with the development of extinction without having any detectable effect on the rats' levels of fear prior to extinction. Injection of FG 7142 also reversed extinction, partially reinstating fear responses that had been extinguished previously. A similar reinstatement of extinguished fear was seen when rats were tested for fear of the cue in a different chamber. The reinstatement produced by FG 7142 and that caused by context shift were not additive: FG 7142 did not increase extinguished fear if rats were tested in the different chamber. Finally, FG 7142 had no detectable effect on the latent inhibition of fear produced by repeatedly presenting the cue alone before conditioning with shock, even though this inhibition, like extinction, was affected by a shift in context. The present findings indicate that GABA transmission at GABA(A) receptors is involved in the inhibition of extinguished fear, and that this effect of GABA is regulated by those cues that constitute the extinction context.

Animals↗

Examination of a backchaining/counterconditioning process during the extinction of conditioned fear.

Two experiments using rat subjects are reported which attempt to delineate the theoretical mechanisms involved in exposure/extinction procedures that are used to eliminate conditioned fear. In Experiment 1, a within-subjects design was employed to study the temporal course of the extinction process by examining the relationship among three separate measures of classically conditioned fear--suppression of an ongoing, operant behavior during the nonreinforced fear-eliciting CS presentation, the time to the first response during this CS presentation, and the time to recover responding following termination of the CS. The results indicated a temporal relationship among these measures, both within and across nonreinforced CS trials, which were considered to reflect a backchaining of a fear-antagonistic response during extinction, and provided an empirical demonstration of an extinction process consistent with Denny's (Behaviour Therapy and Experimental Psychiatry, 7, 315-321, 1976) elicitation/backchaining explanation of extinction. Experiment 2 attempted to manipulate the course of extinction and the temporal relationship among these three fear measures through the paired presentations of the fear CS with a palatable substance (maltose). This procedure produced greater extinction apparently by facilitating the backchaining process. These results are discussed with implications for exposure-based therapies.

Arousal↗

Failure of ACTH to prolong extinction of a conditioned taste aversion in the absence of the testes.

Both corticotropin (ACTH) and testosterone prolong the extinction of a conditioned taste aversion in water-deprived intact male rats. An investigation was made to determine whether ACTH affects extinction in the absence of the testes and also to determine the effect of ACTH on serum testosterone levels. Water-deprived intact males showed prolonged extinction after ACTH injections; water-deprived gonadectomized males and intact females did not. All three of these groups showed elevated testosterone levels after ACTH administration, but testosterone levels were higher in the intact males than in the gonadectomized males or intact females. These results clearly show that in the absence of the testes ACTH is unable to prolong extinction. It is proposed that the increased level of testosterone following ACTH injection in water-deprived intact males is responsible for the prolonged extinction of a conditioned taste aversion. Although testosterone levels may increase in females and castrated males following ACTH injection, the increase is not sufficient to prolong extinction in these water-deprived animals.

Adrenocorticotropic Hormone↗