Comparing the genetic population structure of two species of arctic lemmings: more local differentiation in Lemmus trimucronatus than in Dicrostonyx groenlandicus

Collared and brown lemmings ( Dicrostonyx groenlandicus and Lemmus trimucronatus ) are two largely sympatric and ecologically comparable species of arctic microtine rodents, differing however in some respects which allow us to hypothesise differences in the genetic structure of their populations. Collared lemmings are particularly well adapted to life at high latitude, they occasionally emerge to the surface of the snow and may disperse over larger distances than brown lemmings – possibly even over snow and ice. This should result in more local differentiation among populations of brown lemmings than among populations of collared lemmings. We compared the genetic population structure between the two lemming species in a fragmented landscape with small islands in the central Canadian Arctic using four microsatellite loci and partial mitochondrial control region sequences. Both types of genetic markers showed higher differentiation ( F _ST values) among local populations for brown lemmings than for collared lemmings. We discuss to what extent the observed genetic differences may be explained by differences in dispersal rates in addition to differences in average effective population size.     

Endocrine Correlates of Seasonal Body Mass Dynamics in the Collared Lemming (Dicrostonyx groenlandicus)

Many winter-active temperate rodent species show seasonal, photoperiod-mediatedchanges in body mass that correlate with changes in food availability,gaining mass when food is abundant. The collared lemming (Dicrostonyxgroenlandicus), an arctic species, also undergoes a seasonaltransition in body mass, but one that is temporally out of phasewith that observed in species from lower latitudes; whereastemperate species increase in mass when exposed to relativelylong day lengths, collared lemmings do so under long but decreasingday lengths. This adaptation may be in response tothe timingof peak above-ground biomass in the arctic. Validation of thishypothesis requires a more thorough examination of both collaredlemmings and other arctic species. In the collared lemming, photoperiod-mediated changes in bodymass are correlated with changes in serum concentrations ofprolactin, thyroid hormones, corticosterone, and growth hormone,and pineal concentrations of melatonin. These observations arecompared to those obtained with temperate small mammals, andpossible cause-and-effect relationships between body mass andhormonal parameters are discussed.     

The influence of the antimitotic drug CCNU on the neurosecretion of rat hypothalamo-hypophyseal system

The purpose of this investigation was to evaluate the influence of the antimitotic drug CCNU on the morphology of the hypothalamo-hypophyseal neurosecretory system of rat. Adult Wistar rats were treated intragastrically with 2.5 mg CCNU once a week during 3 consecutive weeks and 5 mg at the end of the 4th week. The brains and hypophyses were fixed in Zenker-formol solution. Paraffin slices were stained with chromhematoxylin to demonstrate neurosecretory material and with cresyl violet. PAS reaction was also performed. The experiment resulted in disturbances of the neurosecretory function of the hypothalamo-hypophyseal system in form of alteration in the content of neurosecretion in the neuronal cytoplasm and processes within supraoptic and paraventricular nuclei as well as in the neurohypophysis. The morphometric measurements showed enlargement of the cell nuclei and cytoplasm volumes in the nucleus supraopticus of hypothalamus.     

Functional and numerical responses of four lemming predators in high arctic Greenland

The high‐arctic tundra ecosystem has the world's simplest vertebrate predator–prey community, with only four predators preying upon one rodent species, the collared lemming (Dicrostonyx groenlandicus). We document the functional and numerical responses of all the four predators in NE Greenland. Using these data, we assess the impact of predation on the dynamics of the collared lemming with a 4 yr cycle and >100‐fold difference between maximum and minimum densities. All predator species feed mostly (>90%) on lemmings when lemming density is >1 ha^?1, but the shapes of the predators’ responses vary greatly. The snowy owl (Nyctea scandiaca) is present and breeds only when lemming densities at snowmelt are >2 ha^?1, giving rise to a step‐like numerical response. The long‐tailed skua (Stercorarius longicaudus) has a type III functional response and shifts from alternate food (mainly berries and insects) to lemmings with increasing lemming density. The skua surpasses all the other predators in summer by its total response. The type III functional response of the Arctic fox (Alopex lagopus) starts to increase at much lower lemming densities than the responses of the avian predators, but it has only a weak numerical response. Finally, the stoat (Mustela erminea) is the most specialized predator and the only one with a clearly delayed numerical response. According to their specific functional and numerical responses, each predator plays a key role at some point of the lemming cycle, but only the stoat has the potential to drive the lemming cycle. Stoat predation is greatly reduced in the winter preceding the lemming peak, and it reaches a maximum in the winter preceding the lowest lemming summer density. Stoat predation appears to maintain low lemming densities for at least two successive years. Our study provides empirical support for the specialist predator hypothesis about small mammal population cycles.     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

Photoperiodic effects on post-weaning growth and food consumption in the collared lemming Dicrostonyx groenlandicus

The effects of long 'summer' (22L:2D) and short 'winter' (2L:22D) photoperiod on post-weaning growth and food consumption of the collared lemming were examined. Growth was described by Gompertz equations. After 10 weeks, lemmings maintained under short photoperiods developed white winter pelage and heavy bifid claws; body mass and estimated asymptotic mass were significantly greater than for lemmings reared under long photoperiod. Sexual dimorphism in body size was observed within treatments, males growing larger than females. There were no significant differences in the overall growth rate constant k as a result of either sex or photoperiod treatment. However, instantaneous growth rates (dW/dt) were significantly higher in 'winter' lemmings. Mass at weaning was a significant determinant of adult mass at 90 days.
Cumulative food consumption at 90 days was not significantly different between photoperiod groups, even though 'winter' lemmings gained more than double the mass of their 'summer' counterparts over the post-weaning period. Increased gut length observed for winter morphs may act as the physiological mechanism promoting greater gross energy efficiency.
The present study suggests that the overwintering strategy of the collared lemming with respect to body mass changes and energy requirements differs considerably from that of other microtines.     

Does lemming winter grazing impact vegetation in the Canadian Arctic?

In low-productivity environments such as the tundra, it has been proposed that regular, multi-annual population cycles of lemmings could be driven by winter food depletion in years of peak abundance. If lemming population dynamics is controlled by food resources, we predict that (1) winter grazing should negatively impact the abundance of food plants, (2) this impact should be proportional to lemming density and (3) high lemming winter grazing pressure should result in reduced plant growth during the following summer. We tested these predictions on Bylot Island, Nunavut, Canada, where two species of lemmings are present: the brown (Lemmus trimucronatus) and collared lemming (Dicrostonyx groenlandicus). We installed 16 exclosures in their preferred wintering habitat (snowbeds) and annually sampled plant biomass inside and outside exclosures at snow melt and at peak growth during the summers of 2009–2012, covering a full population cycle. Winter grazing had no impact on total vascular plant or moss biomass at snow melt in all years. Among plant families, only Caryophyllaceae, which was uncommon, showed a decline. In moss taxa, a negative effect was found on Polytrichum in only 1 year out of three. Overall, plant regrowth during the subsequent summer showed annual variation and tended to be reduced in the 2 years of high lemming abundance. However, this could be a consequence of summer grazing. Overall, the impact of lemming winter grazing on plants was weak and short-lived, even in years of high lemming abundance. Therefore, our results are not consistent with the hypothesis that food depletion during winter was the cause of the lemming decline following peak abundance at our study site. Other factors may limit lemming populations and prevent them from reaching densities high enough to exhaust their food resources.     

The implantation of puromycin into the neurosecretory nuclei of the rat

Summary Following the bilateral implantation of puromycin into the paraventricular nuclei of rats, the neurosecretory cells became atrophic and the amount of aldehyde-fuchsin (AF) positive material in the neural lobe decreased. In these rats, urine excretion and water intake increased remarkably. The supraoptic nuclei of the rats were not affected by this treatment. After the unilateral implantation of puromycin in the paraventricular nucleus, the neurosecretory cells of the implanted side became atrophic, while those of the unimplanted side hypertrophied. The neural lobe contained similar amounts of AF-positive material to those of the control rats with unilateral cholesterol implants. In the rats implanted bilaterally with puromycin immediately above the supraoptic nucleus, the neurosecretory cells of this nucleus contained little or no AF-positive material, and urine excretion and water intake increased greatly. The cells of the paraventricular nucleus remained unchanged in these rats.     

Prolonging the arctic pulse: long-term exploitation of cached eggs by arctic foxes when lemmings are scarce

1. Many ecosystems are characterized by pulses of dramatically higher than normal levels of foods (pulsed resources) to which animals often respond by caching foods for future use. However, the extent to which animals use cached foods and how this varies in relation to fluctuations in other foods is poorly understood in most animals. 2. Arctic foxes Alopex lagopus (L.) cache thousands of eggs annually at large goose colonies where eggs are often superabundant during the nesting period by geese. We estimated the contribution of cached eggs to arctic fox diets in spring and autumn, when geese were not present in the study area, by comparing stable isotope ratios (delta(13)C and delta(15)N) of fox tissues with those of their foods using a multisource mixing model in Program IsoSource. 3. The contribution of cached eggs to arctic fox diets was inversely related to collared lemming Dicrostonyx groenlandicus (Traill) abundance; the contribution of cached eggs to overall fox diets increased from < 28% in years when collared lemmings were abundant to 30-74% in years when collared lemmings were scarce. 4. Further, arctic foxes used cached eggs well into the following spring (almost 1 year after eggs were acquired) - a pattern that differs from that of carnivores generally storing foods for only a few days before consumption. 5. This study showed that long-term use of eggs that were cached when geese were superabundant at the colony in summer varied with fluctuations in collared lemming abundance (a key component in arctic fox diets throughout most of their range) and suggests that cached eggs functioned as a buffer when collared lemmings were scarce.     

Occurrence of the neurosecretory substance during the embryonic development of the guinea pig

Summary The time and place of occurrence of the neurosecretory substance in the hypothalamo-neurohypophysial system of the guinea pig during embryogenesis have been investigated. Use is made of the luminiscence of neurosecretion stained with paraldehydefuchsin when observed in a dark field. It is established that the neurosecretory material occurs first in some cells of the supraoptic nucleus about the 39th–40th day of intrauterine development. In the paraventricular nucleus it is observed about the 44th–45th day. At that time it is seen also in Eminentia mediana and in the neurohypophysis. In the latter, however, it is in a smaller amount than in the areas situated above it. These results are discussed in connection with the transport theory of Bargmann and Scharrer.     

Landscapes of fear or competition? Predation did not alter habitat choice by Arctic rodents

In systems where predation plays a key role in the dynamics of prey populations, such as in Arctic rodents, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, habitat selection by collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings depends on intra- and inter-specific densities, and there has been little agreement on the respective influences of food abundance, predators, and competition for habitat on lemming dynamics. Thus, we investigated whether predation affected selection of sedge-meadow versus upland tundra by collared lemmings in the central Canadian Arctic. We first controlled for the effects of competition on lemming habitat selection. We then searched for an additional signal of predation by comparing habitat selection patterns between 12 control plots and one large grid where lemmings were protected from predators by fencing in 1996 and 1997, but not during 5 subsequent years when we monitored habitat use in the grid as well as in the control plots. Dicrostonyx used upland preferentially over meadows and was more numerous in 1996 and 2011 than in other sample years. Lemmus was also more abundant in 1996 than in subsequent years, but its abundance was too low in the exclosure to assess whether exclusion of predators influenced its habitat selection. Contrary to the effects of competition, predation had a negligible impact on the spatial dynamics of Dicrostonyx, at least during summer. These results suggest that any differences in predation risk between the two habitats have little direct influence on the temporal dynamics of Dicrostonyx even if induced through predator–prey cycles.     

Population dynamics of the collared lemming and the tundra vole at Pearce Point,Northwest Territories,Canada

From 1987 to 1989 we monitored population changes during summer of the collared lemming (Dicrostonyx groenlandicus) and the tundra vole (Microtus oeconomus) at Pearce Point, Northwest Territories, Canada (69° 48 N, 122° 40 W). Populations on four study areas did not cycle but remained at low density (<3/ha) each year and continued at low numbers for the following 3 years (Reid et al. 1995). Lemming numbers often declined throghout the summer in spite of continous reproduction, and population recovery occurred overwinter. Heavy predation losses of radio-collared lemmings occurred each summer and this lemming population may be trapped in a predator-pit. Collared lemmings breed in winter and only because of winter population growth do these populations persist. Tundra vole numbers increased rapidly in most summers but usually declined overwinter. Tundra voles do not seem able to sustain winter reproduction in this extreme environment and this prevents them from reaching high density because of the short summer. Population growth in both these rodents could be prevented by poor food or by predation losses, and landscape patchiness may also help to prevent population growth. For lemmings we do not think that a shortage of shelter or intrinsic limitations could be restricting population increase at Pearce Point. This is the first detailed study of a non-cyclic collared lemming population.     

Habitat selection and the scale of ghostly coexistence among Arctic rodents

Competition between coexisting species existing near their stable equilibrium can be obscured if they occupy separate habitats. Theories of habitat selection promise an ability to reveal the underlying ghost of competition by using isodars to infer the behavioural map of habitat selection. We tested the theory with two years of data on abundance and habitat preference by three Arctic rodent species living at low density along a gradient of wet to dry tundra on Herschel Island in Canada's western Arctic. Generalist brown lemmings exhibited a constant partial preference toward wet tundra whereas specialist collared lemmings and voles occupied the driest and wettest zones respectively. Although both lemming species compete for habitats elsewhere in the Canadian Arctic, isodar analyses suggest that the three species occupy wet and dry habitats independently of one another on Herschel Island. Competition at this large scale may be hidden at low densities, however, if the wet‐dry dichotomy is too coarse. Analyses at a finer subdivision of habitat revealed that these species coexist by using different microhabitats. Collared lemmings shifted their niche towards even drier habitat as the abundance of brown lemmings increased. We were thus able to reveal the ghost of competition lurking at large scales through a more refined analysis at smaller scales of density‐dependent habitat use.     

Effect of progesterone and estrogens on the state of the hypothalamo-hypophyseal neurosecretory system in pregnancy and labor

Morphofunctional state of the hypothalamo-hypophyseal neurosecretory system was studied in rats at late stages of pregnancy. No special differences both in function of the supraoptical and praventricular nuclei, and in the hypophysis of intact and pregnant rats at administration of progesterone were noted. When estrogenes were administered to pregnant animals, a sharp increase in functioning of the supraoptic and especially of the paraventricular nuclei was noted. The greatest activity of the supraoptic nucleus was observed at delivery that could be resulted from a stress reaction to pain.     

Morphofunctional characteristics of neurosecretory cells of the supraoptic nucleus of rats with parathyroprival hypocalcemia]

Humori-positive neurosecretory cells of the supraoptic nucleus of the hypothalamus have been investigated during various time of parathyroprival hypocalcemia after extirpation of the parathyroid glands. Contents of total and ionized calcium, phosphorus in blood serum have been estimated. Volume of nuclei and nucleoli has been measured. In 5 days functional activity of the supraoptic nucleus increases (lightly stained cells predominate, volume of the nuclei and nucleoli increases). In subsequent 15-30 days its activity decreases (increase in amount of dark-stained cells, nucleolar volume decreases). In 60 days there is a tendency to restoration of neurosecretion.     

On the relations between the hypothalamus and the anterior pituitary in Rana temporaria

Summary Dawson (1952) showed that, in Rana, many nerve terminals end round the capillaries of the venous portal system of the median eminence. Our observations not only confirm these facts but show as well that, besides the posterior lobe, the outer zone of the median eminence is also a centre of accumulation of neurosecretory material derived from the pre-optic nucleus. The amount of this neurosecretory material is moreover subject to a certain variation in connection with the seasons and with experimental conditions. The idea is put forward that the outer zone of the median eminence of Rana has to be considered as an analogical organ of the posterior lobe. Indeed, in the median eminence, the same morphological facts can be found, which are advanced for the conception that the posterior lobe is a centre of accumulation and a place of release of neurosecretory material originating from the pre-optic nucleus. On the basis of our observations we presume that the hypothalamo-hypophyseal neurosecretory system of Rana can be divided in two sub-units. System I ends in the posterior lobe; system II in the outer zone of the median eminence. The data of the literature indicate that the two systems also probably exist in the higher vertebrates. From these facts it is obvious that the neurosecretory material is transported from the pre-optic nucleus, via the tract, towards the outer zone of the median eminence.     

Brain – Hatschek's pit relationships in amphioxus species

The vertebrate hypothalamo-hypophyseal neurosecretory system is a complex anatomical device for central nervous control over secretion of pituitary hormones. Since it is present in the most primitive vertebrates, the cyclostomes, it is of interest to look for a possible invertebrate anatomical equivalent, or precursor, for clues as to its evolution. We have found in six species of amphioxus, members of an invertebrate group (cephalochordates), considered to be closest to the vertebrates, that there is a morphologically equivalent neuro-epithelial complex, that in many ways resembles the hypothalamo-hypophyseal system of vertebrates. In the six amphioxus species described here the nervous element is a ventral lobe of the brain, the infundibulum, that extends downward along the right side of the notochord, and ends near the dorsal surface of a Rathke's pouch-like structure known as Hatschek's pit. This part of Hatschek's pit has been found earlier to contain a vertebrate LH-like gonadotropin. Therefore, the infundibulum-Hatschek's pit system of amphioxus may be involved in regulating the seasonal reproductive cycle, and it appears to be a direct homologue of the vertebrate hypothalamo-hypophyseal neurosecretory system functionally as well as morphologically.     

Ultrastructure of the neurosecretory cells of the anterior commissural nucleus of the hypothalamus in rats

The ultrastructure of neurosecretory cells of the anterior commissural nucleus of rat hypothalamus is similar to that of the supraoptic nucleus and of the "magnocellular" part of the paraventricular nucleus. The only difference is a less expressed granular endoplasmatic reticulum and a smaller diameter of elementary neurosecretory granules (80-150 nm in diameter). Such elementary granules are characteristic of neurosecretory terminals located in the external zone of the median eminence. It is suggested that neurosecretory cells of the anterior commissural nucleus project to this neurohemal region.     

The summer decline of the collared lemming, Dicrostonyx groenlandicus, in high arctic Greenland

I studied the population structure and dynamics of the collared lemming, Dicrostonyx groenlandicus Traill, on Traill Island (NE Greenland) in 1998 to 2000. This is the first comprehensive study of a Greenlandic collared lemming population. During this period, population density continuously declined (>100 fold-decrease) from more than 10 ind/ha (peak year 1998) to less than 0.1 ind/ha (low year 2000), with the exception of one period of increase in the winter 1998–99. Several population parameters were correlated with these changes in density. Average body mass was greater in the peak year than in the intermediate year (1999) and greater in August than in July in both years. Growth rates were only positive in 1998 and only for young lemmings. Females had unexpectedly larger home ranges than males and were more mobile than males during the decline phase (1999). Poor recruitment and high predation rates (up to 3.4% per day for females in 1998) were the proximate causes of the summer declines in the study area. A simple calculation shows that even with a maximal rate of increase, this lemming population would not have been able to compensate for the high predation rate, and hence it necessarily faced a predation-induced summer decline during the part of population oscillation studied.