Upland Yedoma taliks are an unpredicted source of atmospheric methane.

Landscape drying associated with permafrost thaw is expected to enhance microbial methane oxidation in arctic soils. Here we show that ice-rich, Yedoma permafrost deposits, comprising a disproportionately large fraction of pan-arctic soil carbon, present an alternate trajectory. Field and laboratory observations indicate that talik (perennially thawed soils in permafrost) development in unsaturated Yedoma uplands leads to unexpectedly large methane emissions (35-78 mg m-2 d-1 summer, 150-180 mg m-2 d-1 winter). Upland Yedoma talik emissions were nearly three times higher annually than northern-wetland emissions on an areal basis. Approximately 70% emissions occurred in winter, when surface-soil freezing abated methanotrophy, enhancing methane escape from the talik. Remote sensing and numerical modeling indicate the potential for widespread upland talik formation across the pan-arctic Yedoma domain during the 21st and 22nd centuries. Contrary to current climate model predictions, these findings imply a positive and much larger permafrost-methane-climate feedback for upland Yedoma.

The Arctic Plant Aboveground Biomass Synthesis Dataset.

Plant biomass is a fundamental ecosystem attribute that is sensitive to rapid climatic changes occurring in the Arctic. Nevertheless, measuring plant biomass in the Arctic is logistically challenging and resource intensive. Lack of accessible field data hinders efforts to understand the amount, composition, distribution, and changes in plant biomass in these northern ecosystems. Here, we present The Arctic plant aboveground biomass synthesis dataset, which includes field measurements of lichen, bryophyte, herb, shrub, and/or tree aboveground biomass (g m-2) on 2,327 sample plots from 636 field sites in seven countries. We created the synthesis dataset by assembling and harmonizing 32 individual datasets. Aboveground biomass was primarily quantified by harvesting sample plots during mid- to late-summer, though tree and often tall shrub biomass were quantified using surveys and allometric models. Each biomass measurement is associated with metadata including sample date, location, method, data source, and other information. This unique dataset can be leveraged to monitor, map, and model plant biomass across the rapidly warming Arctic.

Handling prevents and reverses cognitive deficits induced by sub-chronic phencyclidine in a model for schizophrenia in rats.

Treatments for schizophrenia are not effective in ameliorating cognitive deficits. Therefore, novel therapies are needed to treat cognitive impairments associated with schizophrenia (CIAS), which are modelled in rats through administration of sub-chronic phencyclidine (scPCP). We have previously shown that enrichment via voluntary exercise prevents and reverses impairments in novel object recognition (NOR) in this model. The present study aimed to investigate if handling could prevent delay-induced NOR deficits and prevent and reverse scPCP-induced NOR deficits. Two cohorts of adult female Lister Hooded rats were used. In experiment one, handling (five minutes/day, five days/week for two weeks), took place before scPCP administration (2 mg/kg, i.p. twice-daily for seven days). NOR tests were conducted at two, four, and seven weeks post-handling with a one-minute inter-trial interval (ITI) and at five weeks post-dosing with a six-hour ITI. In experiment two, rats were handled after scPCP administration and tested immediately in the one-minute ITI NOR task and again at two weeks post-handling. In both handling regimens, the scPCP control groups failed to discriminate novelty, conversely the scPCP handled groups significantly discriminated in this task. In the 6 h ITI test, vehicle control and scPCP control failed to discriminate novelty; however, the vehicle handled and scPCP handled groups did significantly discriminate. Handling rats prevented and reversed scPCP-induced deficits and prevented delay-induced NOR deficits. These findings add to evidence that environmental enrichment is a viable treatment for cognitive deficits in rodent tests and models of relevance to schizophrenia, with potential to translate into effective treatments for CIAS.

Cryocampsis: a biophysical freeze-bending response of shrubs and trees under snow loads.

We report a biophysical mechanism, termed cryocampsis (Greek cryo-, cold, + campsis, bending), that helps northern shrubs bend downward under a snow load. Subfreezing temperatures substantially increase the downward bending of cantilever-loaded branches of these shrubs, while allowing them to recover their summer elevation after thawing and becoming unloaded. This is counterintuitive, because biological materials (including branches that show cryocampsis) generally become stiffer when frozen, so should flex less, rather than more, under a given bending load. Cryocampsis involves straining of the cell walls of a branch's xylem (wood), and depends upon the branch being hydrated. Among woody species tested, cryocampsis occurs in almost all Arctic, some boreal, only a few temperate and Mediterranean, and no tropical woody species that we have tested. It helps cold-winter climate shrubs reversibly get, and stay, below the snow surface, sheltering them from winter weather and predation hazards. This should be advantageous, because Arctic shrub bud winter mortality significantly increases if their shoots are forcibly kept above the snow surface. Our observations reveal a physically surprising behavior of biological materials at subfreezing temperatures, and a previously unrecognized mechanism of woody plant adaptation to cold-winter climates. We suggest that cryocampsis' mechanism involves the movement of water between cell wall matrix polymers and cell lumens during freezing, analogous to that of frost-heave in soils or rocks.

Dissociating the effects of distraction and proactive interference on object memory through tests of novelty preference.

Encoding information into memory is sensitive to distraction while retrieving that memory may be compromised by proactive interference from pre-existing memories. These two debilitating effects are common in neuropsychiatric conditions, but modelling them preclinically to date is slow as it requires prolonged operant training. A step change would be the validation of functionally equivalent but fast, simple, high-throughput tasks based on spontaneous behaviour. Here, we show that spontaneous object preference testing meets these requirements in the subchronic phencyclidine rat model for cognitive impairments associated with schizophrenia. Subchronic phencyclidine rats show clear memory sensitivity to distraction in the standard novel object recognition task. However, due to this, standard novel object recognition task cannot assess proactive interference. Therefore, we compared subchronic phencyclidine performance in standard novel object recognition task to that using the continuous novel object recognition task, which offers minimal distraction, allowing disease-relevant memory deficits to be assessed directly. We first determined that subchronic phencyclidine treatment did not affect whisker movements during object exploration. Subchronic phencyclidine rats exhibited the expected distraction standard novel object recognition task effect but had intact performance on the first continuous novel object recognition task trial, effectively dissociating distraction using two novel object recognition task variants. In remaining continuous novel object recognition task trials, the cumulative discrimination index for subchronic phencyclidine rats was above chance throughout, but, importantly, their detection of object novelty was increasingly impaired relative to controls. We attribute this effect to the accumulation of proactive interference. This is the first demonstration that increased sensitivity to distraction and proactive interference, both key cognitive impairments in schizophrenia, can be dissociated in the subchronic phencyclidine rat using two variants of the same fast, simple, spontaneous object memory paradigm.

Same father, same face: Deep learning reveals selection for signaling kinship in a wild primate.

Many animals rely on facial traits to recognize their kin; however, whether these traits have been selected specifically for this function remains unknown. Using deep learning for face recognition, we present the first evidence that interindividual facial resemblance has been selected to signal paternal kinship. Mandrills (Mandrillus sphinx) live in matrilineal societies, in which females spend their entire lives not only with maternal half-sisters (MHS) but also with paternal half-sisters (PHS). We show that PHS have more differentiated social relationships compared to nonkin, suggesting the existence of kin recognition mechanisms. We further demonstrate that facial resemblance increases with genetic relatedness. However, PHS resemble each other visually more than MHS do, despite both kin categories sharing similar degrees of genetic relatedness. This paternally derived facial resemblance among PHS indicates selection to facilitate kin recognition. This study also highlights the potential of artificial intelligence to study phenotypic evolution.

Biogenic volatile organic compound ambient mixing ratios and emission rates in the Alaskan Arctic tundra.

Rapid Arctic warming, a lengthening growing season, and the increasing abundance of biogenic volatile-organic-compound-emitting shrubs are all anticipated to increase atmospheric biogenic volatile organic compounds (BVOCs) in the Arctic atmosphere, with implications for atmospheric oxidation processes and climate feedbacks. Quantifying these changes requires an accurate understanding of the underlying processes driving BVOC emissions in the Arctic. While boreal ecosystems have been widely studied, little attention has been paid to Arctic tundra environments. Here, we report terpenoid (isoprene, monoterpenes, and sesquiterpenes) ambient mixing ratios and emission rates from key dominant vegetation species at Toolik Field Station (TFS; 68°38' N, 149°36' W) in northern Alaska during two back-to-back field campaigns (summers of 2018 and 2019) covering the entire growing season. Isoprene ambient mixing ratios observed at TFS fell within the range of values reported in the Eurasian taiga (0-500 parts per trillion by volume - pptv), while monoterpene and sesquiterpene ambient mixing ratios were respectively close to and below the instrumental quantification limit (~ 2 pptv). Isoprene surface emission rates ranged from 0.2 to 2250 µgC m-2 h-1 (mean of 85 µgC m-2 h-1) and monoterpene emission rates remained, on average, below 1 µgC m-2 h-1 over the course of the study. We further quantified the temperature dependence of isoprene emissions from local vegetation, including Salix spp. (a known isoprene emitter), and compared the results to predictions from the Model of Emissions of Gases and Aerosols from Nature version 2.1 (MEGAN2.1). Our observations suggest a 180 %-215 % emission increase in response to a 3-4°C warming, and the MEGAN2.1 temperature algorithm exhibits a close fit with observations for enclosure temperatures in the 0-30°C range. The data presented here provide a baseline for investigating future changes in the BVOC emission potential of the under-studied Arctic tundra environment.

Variation in δ15 N and δ13 C values of forages for Arctic caribou: effects of location, phenology and simulated digestion.

RATIONALE: The use of stable isotopes for dietary estimates of wildlife assumes that there are consistent differences in isotopic ratios among diet items, and that the differences in these ratios between the diet item and the animal tissues (i.e., fractionation) are predictable. However, variation in isotopic ratios and fractionation of δ13 C and δ15 N values among locations, seasons, and forages are poorly described for arctic herbivores especially migratory species such as caribou (Rangifer tarandus). METHODS: We measured the δ13 C and δ15 N values of seven species of forage growing along a 200-km transect through the range of the Central Arctic caribou herd on the North Slope of Alaska over 2 years. We compared forages available at the beginning (May; n = 175) and the end (n = 157) of the growing season (September). Purified enzymes were used to measure N digestibility and to assess isotopic fractionation in response to nutrient digestibility during simulated digestion. RESULTS: Values for δ13 C declined by 1.38 ‰ with increasing latitude across the transect, and increased by 0.44 ‰ from the beginning to the end of the season. The range of values for δ15 N was greater than that for δ13 C (13.29 vs 5.60 ‰). Differences in values for δ13 C between graminoids (Eriophorum and Carex spp.) and shrubs (Betula and Salix spp.) were small but δ15 N values distinguished graminoids (1.87 ± 1.02 ‰) from shrubs (-2.87 ± 2.93 ‰) consistently across season and latitude. However, undigested residues of forages were enriched in 15 N when the digestibility of N was less than 0.67. CONCLUSIONS: Although δ15 N values can distinguish plant groups in the diet of arctic herbivores, variation in the digestibility of dietary items may need to be considered in applying fractionation values for 15 N to caribou and other herbivores that select highly digestible items (e.g. forbs) as well as heavily defended plants (e.g. woody browse). Published in 2017. This article is a U.S. Government work and is in the public domain in the USA.

Warming and neighbor removal affect white spruce seedling growth differently above and below treeline.

Climate change is expected to be pronounced towards higher latitudes and altitudes. Warming triggers treeline and vegetation shifts, which may aggravate interspecific competition and affect biodiversity. This research tested the effects of a warming climate, habitat type, and neighboring plant competition on the establishment and growth of white spruce (Picea glauca (Moench) Voss) seedlings in a subarctic mountain region. P. glauca seedlings were planted in June 2010 under 4 different treatments (high/control temperatures, with/without competition) in 3 habitats (alpine ridge above treeline/tundra near treeline /forest below treeline habitats). After two growing seasons in 2011, growth, photosynthesis and foliar C and N data were obtained from a total of 156, one-and-a-half year old seedlings that had survived. Elevated temperatures increased growth and photosynthetic rates above and near treeline, but decreased them below treeline. Competition was increased by elevated temperatures in all habitat types. Our results suggest that increasing temperatures will have positive effects on the growth of P. glauca seedlings at the locations where P. glauca is expected to expand its habitat, but increasing temperatures may have negative effects on seedlings growing in mature forests. Due to interspecific competition, possibly belowground competition, the upslope expansion of treelines may not be as fast in the future as it was the last fifty years.

Deletion of α-neurexin II results in autism-related behaviors in mice.

Autism is a common and frequently disabling neurodevelopmental disorder with a strong genetic basis. Human genetic studies have discovered mutations disrupting exons of the NRXN2 gene, which encodes the synaptic adhesion protein α-neurexin II (Nrxn2α), in two unrelated individuals with autism, but a causal link between NRXN2 and the disorder remains unclear. To begin to test the hypothesis that Nrxn2α deficiency contributes to the symptoms of autism, we employed Nrxn2α knockout (KO) mice that genetically model Nrxn2α deficiency in vivo. We report that Nrxn2α KO mice displayed deficits in sociability and social memory when exposed to novel conspecifics. In tests of exploratory activity, Nrxn2α KO mice displayed an anxiety-like phenotype in comparison with wild-type littermates, with thigmotaxis in an open field, less time spent in the open arms of an elevated plus maze, more time spent in the enclosure of an emergence test and less time spent exploring novel objects. However, Nrxn2α KO mice did not exhibit any obvious changes in prepulse inhibition or in passive avoidance learning. Real-time PCR analysis of the frontal cortex and hippocampus revealed significant decreases in the mRNA levels of genes encoding proteins involved in both excitatory and inhibitory transmission. Quantification of protein expression revealed that Munc18-1, encoded by Stxbp1, was significantly decreased in the hippocampus of Nrxn2α KO mice, which is suggestive of deficiencies in presynaptic vesicular release. Our findings demonstrate a causal role for the loss of Nrxn2α in the genesis of autism-related behaviors in mice.

Effects of arctic shrub expansion on biophysical vs. biogeochemical drivers of litter decomposition.

Climate warming in arctic tundra may shift dominant vegetation from graminoids to deciduous shrubs, whose functional traits could, in turn, alter biotic and abiotic controls over biogeochemical cycling of carbon (C) and nitrogen (N). We investigated whether shrub-induced changes in microclimate have stronger effects on litter decomposition and nutrient release than changes in litter quality and quantity. In arctic tundra near Toolik Lake, Alaska, USA, we incubated a common substrate in a snow-addition experiment to test whether snow accumulation around arctic deciduous shrubs altered the environment enough to increase litter decomposition rates. We compared the influence of litter quality on the rate of litter and N loss by decomposing litter from four different plant functional types in a common site. We used aboveground net primary production values and estimated decay constant (k) values from our decomposition experiments to calculate community-weighted mass loss for each site. Snow addition had no effect on decomposition of the common substrate, and the site with the highest abundance of shrubs had the lowest decomposition rates. Species varied in their decomposition rates, with species from the same functional type not always following similar patterns. Community-weighted mass loss was 1.5 times greater in the high shrub site, and only slightly decreased when adjusted for soil environment, suggesting that litter quality and quantity are the primary drivers of community decomposition. Our findings suggest that on a short time scale, the changes in soil environment associated with snow trapping by shrubs are unlikely to influence litter nutrient turnover enough to drive positive snow-shrub feedbacks. The mechanisms driving shrub expansion are more likely to do with shrub-litter feedbacks, where the higher growth rates and N uptake by shrubs allows them to produce more leaves, resulting in a larger litter N pool and faster internal cycling of nutrients.

Phenological response of tundra plants to background climate variation tested using the International Tundra Experiment.

The rapidly warming temperatures in high-latitude and alpine regions have the potential to alter the phenology of Arctic and alpine plants, affecting processes ranging from food webs to ecosystem trace gas fluxes. The International Tundra Experiment (ITEX) was initiated in 1990 to evaluate the effects of expected rapid changes in temperature on tundra plant phenology, growth and community changes using experimental warming. Here, we used the ITEX control data to test the phenological responses to background temperature variation across sites spanning latitudinal and moisture gradients. The dataset overall did not show an advance in phenology; instead, temperature variability during the years sampled and an absence of warming at some sites resulted in mixed responses. Phenological transitions of high Arctic plants clearly occurred at lower heat sum thresholds than those of low Arctic and alpine plants. However, sensitivity to temperature change was similar among plants from the different climate zones. Plants of different communities and growth forms differed for some phenological responses. Heat sums associated with flowering and greening appear to have increased over time. These results point to a complex suite of changes in plant communities and ecosystem function in high latitudes and elevations as the climate warms.

The response of Arctic vegetation and soils following an unusually severe tundra fire.

Fire causes dramatic short-term changes in vegetation and ecosystem function, and may promote rapid vegetation change by creating recruitment opportunities. Climate warming likely will increase the frequency of wildfire in the Arctic, where it is not common now. In 2007, the unusually severe Anaktuvuk River fire burned 1039 km(2) of tundra on Alaska's North Slope. Four years later, we harvested plant biomass and soils across a gradient of burn severity, to assess recovery. In burned areas, above-ground net primary productivity of vascular plants equalled that in unburned areas, though total live biomass was less. Graminoid biomass had recovered to unburned levels, but shrubs had not. Virtually all vascular plant biomass had resprouted from surviving underground parts; no non-native species were seen. However, bryophytes were mostly disturbance-adapted species, and non-vascular biomass had recovered less than vascular plant biomass. Soil nitrogen availability did not differ between burned and unburned sites. Graminoids showed allocation changes consistent with nitrogen stress. These patterns are similar to those seen following other, smaller tundra fires. Soil nitrogen limitation and the persistence of resprouters will likely lead to recovery of mixed shrub-sedge tussock tundra, unless permafrost thaws, as climate warms, more extensively than has yet occurred.

Carbon loss from an unprecedented Arctic tundra wildfire.

Arctic tundra soils store large amounts of carbon (C) in organic soil layers hundreds to thousands of years old that insulate, and in some cases maintain, permafrost soils. Fire has been largely absent from most of this biome since the early Holocene epoch, but its frequency and extent are increasing, probably in response to climate warming. The effect of fires on the C balance of tundra landscapes, however, remains largely unknown. The Anaktuvuk River fire in 2007 burned 1,039 square kilometres of Alaska's Arctic slope, making it the largest fire on record for the tundra biome and doubling the cumulative area burned since 1950 (ref. 5). Here we report that tundra ecosystems lost 2,016 ± 435 g C m(-2) in the fire, an amount two orders of magnitude larger than annual net C exchange in undisturbed tundra. Sixty per cent of this C loss was from soil organic matter, and radiocarbon dating of residual soil layers revealed that the maximum age of soil C lost was 50 years. Scaled to the entire burned area, the fire released approximately 2.1 teragrams of C to the atmosphere, an amount similar in magnitude to the annual net C sink for the entire Arctic tundra biome averaged over the last quarter of the twentieth century. The magnitude of ecosystem C lost by fire, relative to both ecosystem and biome-scale fluxes, demonstrates that a climate-driven increase in tundra fire disturbance may represent a positive feedback, potentially offsetting Arctic greening and influencing the net C balance of the tundra biome.

Linking functional diversity and social actor strategies in a framework for interdisciplinary analysis of nature's benefits to society.

The crucial role of biodiversity in the links between ecosystems and societies has been repeatedly highlighted both as source of wellbeing and as a target of human actions, but not all aspects of biodiversity are equally important to different ecosystem services. Similarly, different social actors have different perceptions of and access to ecosystem services, and therefore, they have different wants and capacities to select directly or indirectly for particular biodiversity and ecosystem characteristics. Their choices feed back onto the ecosystem services provided to all parties involved and in turn, affect future decisions. Despite this recognition, the research communities addressing biodiversity, ecosystem services, and human outcomes have yet to develop frameworks that adequately treat the multiple dimensions and interactions in the relationship. Here, we present an interdisciplinary framework for the analysis of relationships between functional diversity, ecosystem services, and human actions that is applicable to specific social environmental systems at local scales. We connect the mechanistic understanding of the ecological role of diversity with its social relevance: ecosystem services. The framework permits connections between functional diversity components and priorities of social actors using land use decisions and ecosystem services as the main links between these ecological and social components. We propose a matrix-based method that provides a transparent and flexible platform for quantifying and integrating social and ecological information and negotiating potentially conflicting land uses among multiple social actors. We illustrate the applicability of our framework by way of land use examples from temperate to subtropical South America, an area of rapid social and ecological change.

Schizophrenia-related endophenotypes in heterozygous neuregulin-1 'knockout' mice.

Neuregulin-1 (NRG1) has been shown to play a role in glutamatergic neurotransmission and is a risk gene for schizophrenia, in which there is evidence for hypoglutamatergic function. Sensitivity to the behavioural effects of the psychotomimetic N-methyl-D-aspartate receptor antagonists MK-801 and phencyclidine (PCP) was examined in mutant mice with heterozygous deletion of NRG1. Social behaviour (sociability, social novelty preference and dyadic interaction), together with exploratory activity, was assessed following acute or subchronic administration of MK-801 (0.1 and 0.2 mg/kg) or PCP (5 mg/kg). In untreated NRG1 mutants, levels of glutamate, N-acetylaspartate and GABA were determined using high-performance liquid chromatography and regional brain volumes were assessed using magnetic resonance imaging at 7T. NRG1 mutants, particularly males, displayed decreased responsivity to the locomotor-activating effects of acute PCP. Subchronic MK-801 and PCP disrupted sociability and social novelty preference in mutants and wildtypes and reversed the increase in both exploratory activity and social dominance-related behaviours observed in vehicle-treated mutants. No phenotypic differences were demonstrated in N-acetylaspartate, glutamate or GABA levels. The total ventricular and olfactory bulb volume was decreased in mutants. These data indicate a subtle role for NRG1 in modulating several schizophrenia-relevant processes including the effects of psychotomimetic N-methyl-D-aspartate receptor antagonists.

Ecological dynamics across the Arctic associated with recent climate change.

At the close of the Fourth International Polar Year, we take stock of the ecological consequences of recent climate change in the Arctic, focusing on effects at population, community, and ecosystem scales. Despite the buffering effect of landscape heterogeneity, Arctic ecosystems and the trophic relationships that structure them have been severely perturbed. These rapid changes may be a bellwether of changes to come at lower latitudes and have the potential to affect ecosystem services related to natural resources, food production, climate regulation, and cultural integrity. We highlight areas of ecological research that deserve priority as the Arctic continues to warm.

Plant functional types do not predict biomass responses to removal and fertilization in Alaskan tussock tundra.

Plant communities in natural ecosystems are changing and species are being lost due to anthropogenic impacts including global warming and increasing nitrogen (N) deposition. We removed dominant species, combinations of species and entire functional types from Alaskan tussock tundra, in the presence and absence of fertilization, to examine the effects of non-random species loss on plant interactions and ecosystem functioning.After 6 years, growth of remaining species had compensated for biomass loss due to removal in all treatments except the combined removal of moss, Betula nana and Ledum palustre (MBL), which removed the most biomass. Total vascular plant production returned to control levels in all removal treatments, including MBL. Inorganic soil nutrient availability, as indexed by resins, returned to control levels in all unfertilized removal treatments, except MBL.Although biomass compensation occurred, the species that provided most of the compensating biomass in any given treatment were not from the same functional type (growth form) as the removed species. This provides empirical evidence that functional types based on effect traits are not the same as functional types based on response to perturbation. Calculations based on redistributing N from the removed species to the remaining species suggested that dominant species from other functional types contributed most of the compensatory biomass.Fertilization did not increase total plant community biomass, because increases in graminoid and deciduous shrub biomass were offset by decreases in evergreen shrub, moss and lichen biomass. Fertilization greatly increased inorganic soil nutrient availability.In fertilized removal treatments, deciduous shrubs and graminoids grew more than expected based on their performance in the fertilized intact community, while evergreen shrubs, mosses and lichens all grew less than expected. Deciduous shrubs performed better than graminoids when B. nana was present, but not when it had been removed.Synthesis. Terrestrial ecosystem response to warmer temperatures and greater nutrient availability in the Arctic may result in vegetative stable-states dominated by either deciduous shrubs or graminoids. The current relative abundance of these dominant growth forms may serve as a predictor for future vegetation composition.