Equipped for Life in the Boreal Forest: The Role of the Stress Axis in Mammals

The hypothalamic-pituitary-adrenal axis (stress axis) plays a central role in equipping mammals to succeed in the challenging environment of the boreal forest. Over the last 20 years, we have tackled a broad range of topics to understand how the stress axis functions in four key herbivore species. The central challenge for snowshoe hares is coping with their predators, whereas for the others, it is primarily coping with each other (especially during reproduction) and with their physical environment. Hares are severely stressed by their predators during the population decline. The predator threat causes major changes in the stress axis of hares and reduces their reproduction; in addition, acting through maternal programming, it is the most plausible explanation for the extended period of low numbers following the population decline. Arctic ground squirrel males have an intense breeding season for two to three weeks in early spring, after which many of them die. The functioning of their stress axis changes markedly and is key in meeting their energy demands during this period. In contrast, red-backed vole males, though also short-lived, breed repeatedly only in the summer of their life, and their stress axis shows no change in function. However, their reproductive effort negatively affects their long-term survival. Territorial red squirrels experience marked interannual fluctuations in their major food source (white spruce seed), resulting in major fluctuations in their densities and consequently in the intensity of territorial competition. Changes in the densities of red squirrels also alter maternal stress hormone levels, inducing adaptive plasticity in offspring postnatal growth rates that prepares offspring for the environment they will encounter at independence. To survive winter, red squirrels need to defend their territories year-round, and the basis of this defense appears to be adrenal dehydroepiandrosterone, which has the benefits, but not the costs, of gonadal steroids. Arctic ground squirrels survive winter by hibernating in deeply frozen ground. Unlike all other hibernators, they have evolved a unique adaptation: high levels of adrenal androgens in summer to accumulate protein reserves that they then burn in winter. With a rapidly changing climate, the stress axis will play a key role in permitting northern animals to adapt, but the linkages between the changes in the abiotic and biotic components of the boreal forest and the phenotypic plasticity in the stress response of its inhabitants are not well understood for these or any other herbivore species.L’axe hypothalamo-hypophyso-surrénalien (l’axe du stress) joue un rôle central pour aider les mammifères à réussir dans l’environnement difficile de la forêt boréale. Ces 20 dernières années, nous nous sommes penchés sur une vaste gamme de sujets afin de comprendre comment fonctionne l’axe du stress chez quatre grandes espèces herbivores. Pour le lièvre d’Amérique, le défi central consiste à faire face à ses prédateurs, tandis que pour les autres espèces, ce défi consiste à se faire face mutuellement (surtout pendant la reproduction) de même qu’à faire face à leur environnement physique. Les lièvres subissent beaucoup de stress de la part de leurs prédateurs pendant la diminution de la population. La menace des prédateurs est la cause de changements majeurs sur l’axe du stress des lièvres, ce qui a pour effet de réduire leur reproduction. De plus, en raison de leur programmation maternelle, il s’agit de l’explication la plus plausible justifiant la période prolongée de leur faible nombre suivant la diminution de la population. Le spermophile arctique mâle a une période de reproduction intense pendant deux à trois semaines au début du printemps et après cela, un grand nombre d’entre eux meurent. Le fonctionnement de son axe de stress change de façon marquée, ce qui est essentiel à sa demande en énergie pendant cette période. Par contraste, le campagnol à dos roux mâle, même s’il ne vit également pas longtemps, se reproduit à répétition seulement pendant l’été de sa vie, et le fonctionnement de son axe de stress ne montre aucun changement. Cependant, ses efforts de reproduction ont des incidences négatives sur sa survie à long terme. Pour sa part, la principale source d’alimentation (les graines d’épinette blanche) de l’écureuil roux territorial connaît des fluctuations interannuelles marquées, ce qui se traduit par une fluctuation majeure en matière de densité de cette espèce animale et, par conséquent, en matière d’intensité de la concurrence pour le territoire. Les changements de densité d’écureuils roux exercent également une influence sur les taux d’hormone maternelle de stress, ce qui donne lieu à une plasticité adaptative des taux de croissance postnatale de la progéniture qui prépare la progéniture pour faire face à l’environnement dans lequel ils évolueront au stade de l’indépendance. Pour survivre à l’hiver, l’écureuil roux doit défendre son territoire à l’année et pour y parvenir, il se sert de la déhydroépiandrostérone surrénalienne, qui comporte les avantages des stéroïdes gonades, sans les coûts. Le spermophile arctique survit à l’hiver en hibernant dans le sol gelé en profondeur. Contrairement à tous les autres hibernateurs, il s’est développé une adaptation unique en son genre, soit des taux élevés d’androgènes surrénaliens en été qui lui permettent d’accumuler les réserves de protéines qu’il brûle ensuite pendant l’hiver. À la lumière du changement climatique rapide, l’axe de stress jouera un rôle-clé pour permettre aux animaux du Nord de s’adapter, mais les liens entre les changements des composantes abiotiques et biotiques de la forêt boréale et la plasticité phénotypique de la réaction de stress de ses habitants ne sont pas bien compris dans le cas de ces espèces herbivores ou de toute autre espèce herbivore

Factors Influencing Common Raven Occurrence and Density Across Cold-Desert Sagebrush Ecosystems of the Southwestern U.S.

Common ravens (Corvus corax) are a predator of eggs and chicks of numerous species including greater sage-grouse (Centrocercus urophasianus). Raven abundance and distribution is increasing within sagebrush ecosystems as a result of anthropogenic resource subsidies. Despite concerns about subsequent predation pressure on sage-grouse, broad-scale spatial information about raven populations remains lacking. We used hierarchical occupancy and distance sampling models to map raven density and distribution in response to natural and anthropogenic landscape covariates using \u3e15,000 point count surveys occurring within the Great Basin region since 2007. Anthropogenic factors contributing to greater raven occurrence included increased road density, presence of transmission lines, agricultural activity, and presence of roadside rest areas. Natural landscape characteristics included lower elevations with greener vegetation (NDVI), greater stream and habitat edge densities, and lower percentages of big sagebrush (A. tridentate spp.). Many of these same environmental factors influenced spatial variation in raven density, although the effects varied by field site. Both raven occurrence and density tended to increase in valleys with networks of agricultural fields, ranches, roads, and distribution lines. These features likely subsidize local raven populations, which then move into more remote shrubland environments with negative consequences for sage-grouse populations. We used the relationships identified in our model to make predictions of raven density and distribution across the Great Basin landscape. We show how these model outputs can be used to guide management decisions where raven distributions overlap with breeding sage-grouse concentration areas. Findings are preliminary and provided for timely best science

Effects of Common Ravens on Greater Sage-Grouse in the Great Basin, Region, USA

Anthropogenic modification to ecosystems can result in the redistribution of species at higher trophic levels. Humans have re-organized predator-prey dynamics, namely by removing top predators and subsidizing more generalist mesocarnivore species. As a result, some mid-level predator species have increased in abundance and distribution, often to the detriment of lower-level species that are not adapted to increased predation rates. One example of a native avian predator that has experienced population increase following increased anthropogenic subsidization is the common raven (Corvus corax; hereafter, raven).The raven is an ubiquitous predator within sagebrush ecosystems in the western U.S.,and may contribute to suppressed population growth in greater sage-grouse (Centrocercus urophasianus) through disruptions to lekking behavior and top-down influences on nest success and recruitment. Ravens have expanded in distribution and abundance, in large part due to increased resource subsidies from human infrastructure and land use activities. Concurrently, some sage-grouse populations appear to be in decline where habitat conditions should be promoting species persistence. Using long-term monitoring data on sage-grouse and ravens in the northern Great Basin region, we show that ravens disrupt sage-grouse lekking behavior, increased raven density is associated with reduced sage-grouse nest success, and that negative trends in lek counts may be related to elevated raven occurrence and density. Taken together, these results suggest the need to address a growing problem, as ravens continue to expand their distribution, facilitated by anthropogenic subsidies. These findings are preliminary and provided to meet the need for timely best science

A Rapid Assessment Function to Estimate Common Raven Population Densities: Implications for Targeted Management

Common raven (Corvus corax; raven) populations have increased over the past 5 decades within the western United States. Raven population increases have been largely attributed to growing resource subsidies from expansion of human enterprise. Concomitantly, managers are becoming increasingly concerned about elevated adverse effects on multiple sensitive prey species, damage to livestock and agriculture, and human safety. Managers could benefit from a rapid but reliable method to estimate raven densities across spatiotemporal scales to monitor raven populations more efficiently and inform targeted and adaptive management frameworks. However, obtaining estimates of raven density is data- and resource-intensive, which renders monitoring within an adaptive framework unrealistic. To address this need, we developed a rapid survey protocol for resource managers to estimate site-level density based on the average number of ravens per survey. Specifically, we first estimated raven densities at numerous field sites with robust distance sampling procedures and then used regression to investigate the relationship between those density estimates and the number of ravens per survey, which revealed a strong correlation (R2 = 0.86). For management application, we provide access to R function software through a web-based interface to estimate density using number of ravens per survey, which we refer to as a Rapid Assessment Function (RAF). Then, using a simulation analysis of data from sites with abundant surveys and the RAF, we estimated raven density based on different numbers of surveys to help inform how many surveys are needed to achieve reliable estimates within this rapid assessment. While more robust procedures of distance sampling are the preferred methods for estimating raven densities from count surveys, the RAF tool presented herein provides a reliable approximation for informing management decisions when managers are faced with resource and small sample size constraints

Spatial Modeling of Common Raven Density and Occurrence Helps Guide Landscape Management Within Great Basin Sagebrush Ecosystems

Common ravens (Corvus corax; ravens) are a behaviorally flexible nest predator of several avian species, including species of conservation concern. Movement patterns based on life history phases, particularly territoriality of breeding birds and transiency of nonbreeding birds, are thought to influence the frequency and efficacy of nest predation. As such, predicting where on the landscape territorial resident and non-territorial transient birds may be found in relation to the distribution of sensitive prey is of increasing importance to managers and conservationists. From 2007 to 2019, we conducted raven point count surveys between mid-March and mid-September across 43 different field sites representing typical sagebrush (Artemisia spp.) ecosystems of the Great Basin, USA. The surveys conducted during 2007–2016 were used in previously published maps of raven occurrence and density. Here, we examined the relationship between occurrence and density of ravens using spatially explicit predictions from 2 previously published studies and differentiate areas occupied by higher concentrations of resident ravens as opposed to transients. Surveys conducted during 2017–2019 were subsequently used to evaluate the predicted trends from our analytical approach. Specifically, we used residuals from a generalized linear regression to establish the relationship between occurrence and density, which ultimately resulted in a spatially explicit categorical map that identifies areas of resident versus transient ravens. We evaluated mapped categories using independently collected observed raven group sizes from the 2017–2019 survey data, as well as an independent dataset of global positioning system locations of resident and transient individuals monitored during 2019–2020. We observed moderate agreement between the mapped categories and independent datasets for both evaluation approaches. Our map provides broad inference about spatial variation in potential predation risk from ravens for species such as greater sage-grouse (Centrocercus urophasianus) and can be used as a valuable spatial layer for decision support tools aimed at guiding raven management decisions and, ultimately, improving survival and reproduction of sensitive prey within the Great Basin

Inherited electrophoretic variants detected in a Japanese population with two-dimensional gels of erythrocyte lysates

Genetic variation has been studied in erythrocyte lysates from 100 Japanese children and their parents by means of two-dimensional polyacrylamide gel electrophoresis. Fifty-five polypeptides selected without respect to variability were considered suitable for scoring. Genetic variation was encountered in 14 of these polypeptides. Family data show that the segregation of 13 variants is consistent with an autosomal codominant mode of inheritance; the remaining variant exhibits a sex-linked mode of inheritance. Of 8 presumably identical polypeptides found variable in Japanese and/or Caucasians, differences in the occurrence or allele frequencies of polymorphisms were recognized for four. Contrary to the experience of some investigators, the amount of variation and the ethnic differences we are encountering indicate that two-dimensional polyacrylamide gel electrophoresis is a sensitive tool for the study of genetic events.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/65555/1/j.1469-1809.1986.tb01753.x.pd

Efficacy of Manipulating Reproduction of Common Ravens to Conserve Sensitive Prey Species: Three Case Studies

Expansion of human enterprise across western North America has resulted in an increase in availability of anthropogenic resource subsidies for generalist species. This has led to increases in generalists’ population numbers across landscapes that were previously less suitable for their current demographic rates. Of particular concern are growing populations of common ravens (Corvus corax; ravens), because predation by ravens is linked to population declines of sensitive species. Ecosystem managers seek management options for mitigating the adverse effects of raven predation where unsustainable predator–prey conflicts exist. We present 3 case studies examining how manipulating reproductive success of ravens influences demographic rates of 2 sensitive prey species. Two case studies examine impacts of removing raven nests or oiling raven eggs on nest survival of greater sage-grouse (Centrocercus urophasianus; sage-grouse) within Wyoming and the Great Basin of California and Nevada, USA, respectively. The third case study uses Mojave desert tortoise (Gopherus agassizii; tortoise) decoys to examine effects of oiling raven eggs on depredation rates of juvenile tortoises in the Mojave Desert in California. Initial trial years from all 3 case studies were consistent in finding improved vital rates associated with the application of strategies for reducing reproductive success of ravens. Specifically, removal of raven nests resulted in increased nest survival of sage-grouse within treatment areas where predation by ravens was the primary cause of nest failure. In addition, nest survival of sage-grouse and survival of juvenile tortoise decoys was higher following a treatment of oiling the eggs of ravens in their nests at 2 sites within the Great Basin and 4 tortoise conservation areas in the Mojave Desert in California. Along with specialized technologies that can make techniques such as egg-oiling more feasible, these findings support these management practices as important tools for managing ravens, especially in areas where breeding ravens have negative impacts on sensitive prey species

An HDG Method for Dirichlet Boundary Control of Convection Dominated Diffusion PDE

We first propose a hybridizable discontinuous Galerkin (HDG) method to approximate the solution of a \emph{convection dominated} Dirichlet boundary control problem. Dirichlet boundary control problems and convection dominated problems are each very challenging numerically due to solutions with low regularity and sharp layers, respectively. Although there are some numerical analysis works in the literature on \emph{diffusion dominated} convection diffusion Dirichlet boundary control problems, we are not aware of any existing numerical analysis works for convection dominated boundary control problems. Moreover, the existing numerical analysis techniques for convection dominated PDEs are not directly applicable for the Dirichlet boundary control problem because of the low regularity solutions. In this work, we obtain an optimal a priori error estimate for the control under some conditions on the domain and the desired state. We also present some numerical experiments to illustrate the performance of the HDG method for convection dominated Dirichlet boundary control problems

Dissection of the Transformation of Primary Human Hematopoietic Cells by the Oncogene NUP98-HOXA9

NUP98-HOXA9 is the prototype of a group of oncoproteins associated with acute myeloid leukemia. It consists of an N-terminal portion of NUP98 fused to the homeodomain of HOXA9 and is believed to act as an aberrant transcription factor that binds DNA through the homeodomain. Here we show that NUP98-HOXA9 can regulate transcription without binding to DNA. In order to determine the relative contributions of the NUP98 and HOXA9 portions to the transforming ability of NUP98-HOXA9, the effects of NUP98-HOXA9 on primary human CD34+ cells were dissected and compared to those of wild-type HOXA9. In contrast to previous findings in mouse cells, HOXA9 had only mild effects on the differentiation and proliferation of primary human hematopoietic cells. The ability of NUP98-HOXA9 to disrupt the differentiation of primary human CD34+ cells was found to depend primarily on the NUP98 portion, whereas induction of long-term proliferation required both the NUP98 moiety and an intact homeodomain. Using oligonucleotide microarrays in primary human CD34+ cells, a group of genes was identified whose dysregulation by NUP98-HOXA9 is attributable primarily to the NUP98 portion. These include RAP1A, HEY1, and PTGS2 (COX-2). Their functions may reflect the contribution of the NUP98 moiety of NUP98-HOXA9 to leukemic transformation. Taken together, these results suggest that the effects of NUP98-HOXA9 on gene transcription and cell transformation are mediated by at least two distinct mechanisms: one that involves promoter binding through the homeodomain with direct transcriptional activation, and another that depends predominantly on the NUP98 moiety and does not involve direct DNA binding