A previous study reported that a 400-mg dose of medroxyprogesterone acetate (MPA) reduced male reindeer aggression and blocked development of secondary sexual characteristics but did not completely impair fertility. Here we have repeated that protocol in two separate trials. In 2017, tissues and blood samples, collected from MPA and control (CTL) reindeer bulls, euthanized at 30 and 60 d post-treatment were used to evaluate testes histology and morphometrics, cfos activity in the brain and androgen levels. While testes weight tended to decline from August to September in both groups, indices of spermatogenesis remained high. By September, indices of spermatogenesis were declining in both groups with sperm density lower (P = 0.05) in MPA compared to CTL bulls. Aug CTL bulls had the highest concentrations of androstenedione (A4) (P = 0.009) and testosterone (T) (P = 0.08), whereas these androgens were baseline in Aug MPA bulls. By September, A4 and T levels in CTL bulls declined to levels measured in MPA bulls. Cfos activity had a greater number (P = 0.02) of cfos positive neurons in the central amygdala in MPA compared to CTL bulls, suggesting a heightened fear response among the MPA bulls. In the second trial (2019), MPA-treated bulls, with (E, n = 4) and without (IE, n = 4) breeding experience, were blood sampled at key points from July through September when they were put in individual harems with estrous-synchronized cows. Concentrations of T were greatest (P < 0.001) among E bulls prior to MPA treatment but 1 mo after treatment, both T and A4 were baseline in all eight reindeer. Semen collected by electroejaculation at 60 d post-MPA treatment revealed only minor differences in sperm abnormalities between E and IE bulls using both fresh and frozen/thawed semen. Only three bulls (2 E and 1 IE) sired offspring. Breeding success was not related to previous breeding experience, body weight, or bull age. The failure of some MPA bulls to breed appears to be a behavioral, not a physiological, limitation. Limited application of MPA is clearly a useful tool for managing rut-aggression in non-breeding reindeer. However, the possibility that semen could be collected from MPA-treated bulls using restraint and mild sedation rather than general anesthesia should be investigated. This could improve the quality of semen collection while enhancing the safety of both handlers and animals.

Muskoxen (Ovibos moschatus), a taxonomically unique Arctic species, are increasingly exposed to climate and other anthropogenic changes. It is critical to develop and validate reliable tools to monitor their physiological stress response in order to assess the impacts of these changes. Here, we measured fecal glucocorticoid metabolite (FGM) levels in response to the administration of adrenocorticotropic hormone (ACTH) in the winter (1 IU/kg) and summer (2 IU/kg) using two enzyme immunoassays, one targeting primarily cortisol and the other targeting primarily corticosterone. Fecal cortisol levels varied substantially within and among individuals, and none of the animals in either challenge showed an increase in fecal cortisol following the injection of ACTH. By contrast, two of six (winter) and two of five (summer) muskoxen showed a clear response in fecal corticosterone levels (i.e., maximal percentage increase as compared to time 0 levels > 100%). Increases in fecal corticosterone post-ACTH injection occurred earlier and were of shorter duration in the summer than in the winter and fecal corticosterone levels were, in general, lower during the summer. These seasonal differences in FGM responses may be related to the use of different individuals (i.e., influence of sex, age, social status, etc.) and to seasonal variations in the metabolism and excretion of glucocorticoids, intestinal transit time, voluntary food intake, and fecal output and moisture content. Results from this study support using FGMs as a biomarker of hypothalamic-pituitary-adrenal axis activity in muskoxen, advance our understanding of the physiological adaptations of mammals living in highly seasonal and extreme environments such as the Arctic, and emphasize the importance of considering seasonality in other species when interpreting FGM levels.

Muskoxen (Ovibos moschatus) are increasingly exposed to a broad diversity of stressors in their rapidly changing Arctic environment. There is an urgent need to develop validated tools to monitor the impact of these stressors on the hypothalamic-pituitary-adrenal (HPA) axis activity of muskoxen to help inform conservation actions. Here, we evaluated whether muskox qiviut (dense wooly undercoat) cortisol accurately reflects changes in HPA axis activity. Two repeated pharmacological challenges, involving weekly administrations of saline (control group) or adrenocorticotropic hormone (ACTH) during five consecutive weeks, were done on captive muskoxen, in winter (no hair growth) and summer (maximum hair growth). Pre-challenge qiviut cortisol levels were significantly higher in the shoulder than in the neck, but neither differed from rump concentrations. Qiviut cortisol levels significantly increased (p < 0.001) in response to the administration of ACTH during the hair growth phase, but not in the absence of growth (p = 0.84). Cortisol levels in the qiviut segment grown during the summer challenge increased significantly over a six-month period in the ACTH-injected muskoxen with a similar trend occurring in the control animals. Finally, cortisol levels in shed qiviut were significantly higher and not correlated to those of fully grown qiviut shaved three months earlier. Our results show that cortisol is deposited in qiviut during its growth and that qiviut cortisol can thus be used as an integrated measure of HPA axis activity over the period of the hair's growth. Differences in qiviut cortisol across body regions, significant differences in qiviut segments over time, and differences between shed qiviut versus unshed qiviut, highlight the importance of consistent design and methodology for sample collection and analyses in order to account for sources of variation when using qiviut cortisol as a biomarker of HPA axis activity in muskoxen.

Reindeer (Rangifer tarandus tarundus) are the only cervids indigenous to the arctic environment. In Alaska, reindeer are a recognized agricultural species and an economic mainstay for many native populations. Traditionally raised in extensive free-ranging systems, a recent trend toward intensive farming requires a more in-depth knowledge of reproductive management. Reported gestation length in reindeer varies, ranging from 198 to 229 d in studies performed at the University of Alaska Fairbanks. A switchback study that manipulated only breeding date demonstrated a mean increase in gestation length of 8.5 d among females bred early in the season. The negative correlation between conception date and gestation length is consistent with reindeer research at other locations and reports of variable gestation length in a growing number of domestic and non-domestic species. This paper reviews the phenomenon in reindeer and discusses some of the factors known to affect gestation length as well as possible areas for future research.

Energy is a critical driver of animal life-history traits and Darwinian fitness, with direct implications for population dynamics. In the Arctic, extreme seasonal climatic conditions dictate energy flow and physiology of resident species. Consequently, vital rates of animal populations in the Arctic are tightly linked to climatic factors influencing food availability and quality. The muskox (Ovibos moschatus) is a capital breeder and the largest herbivore in the Arctic. Seasonal changes in energy intake influence body reserve dynamics of this species, impacting inter-annual and spatial variations in survival and recruitment. Yet, little is known about the full lifecycle bioenergetics of this and other Arctic species, and the impact of extreme seasonal conditions on life history traits have never been quantified. By integrating energetic and physiological processes throughout the full lifecycle of individuals, we developed an individual-based model using dynamic energy budget theory (DEB-IBM) of the muskox that covers all aspects of mammalian life-history. We parameterized a baseline muskox model using data from a well-studied captive research herd, and then explored its application to a wild population in the high Arctic to assess the life-history implications of seasonal food availability. Our baseline DEB-IBM, which for the first time explicitly characterized fetal development using DEB theory, successfully predicted life-history traits as well as patterns of growth, body condition, feeding, metabolic rates, and reproduction of captive muskoxen. Adaptation of the model to the wild population required re-parameterization of only three variables that controlled seasonal food availability and upregulated assimilation. The wild model successfully captured inter-annual costs of reproduction, such that lactating females were in poorer body condition than females without calves. Arctic seasonal food limitations reduced fecundity in wild muskoxen by 42% compared to annual reproduction in well-fed captive individuals. We have developed a mechanistic energy-based framework to study life-history processes of muskoxen but the model is broadly applicable also for other mammals in the Arctic. Our study quantifies and highlights the strong interplay between energetics and individual fitness components, opening the door to future applications of the model in assessments of population dynamics and climate change impacts.

Reindeer bulls are difficult to manage and dangerous to handlers during the rutting period. Progesterone agonists have been used anecdotally in the field to favorably influence behavior, but effects on reproductive signaling have not been determined. The objective of this study was to determine the effects of Depo-Provera (medroxyprogesterone acetate) on neural activity in the amygdala of reindeer bulls in the early (n = 4) and full (n = 4) rut. Treated bulls (n = 4) were injected with a single injection of Depo-Provera (400 mg i.m.) approximately 2 wk before rut was initiated. Control bulls were untreated. Bulls were exsanguinated and brains collected. Neural activity in the amygdala was determined using c-fos immunohistochemistry. Neural activity did not differ by treatment (P ≥ 0.5), collection period (P ≥ 0.5), or their interaction (P ≥ 0.3) in the medial and cortical amygdala nuclei. A treatment × time interaction (P = 0.009) was observed in the central amygdala. The amygdala nuclei are interconnected allowing for integration of sensory stimuli with a direct connection between the medial amygdala and the olfactory bulb. The central amygdala is responsible for alerting, fear, and initiating a state of arousal towards nonspecific stimuli in the surrounding environment. In wildlife, the central amygdala has a role in recognizing threats in the environment such as predators. During the rut, bulls normally have a decreased sense of fear and elevated aggressive behavior with Depo-Provera treatment seemly able to diminish that aggression. Although it is unlikely that this observed change in neural activity fully explains the decreased aggressive behavior noted in bulls treated with Depo-Provera, neural networks of aggression include the amygdala. It is possible that further changes in c-fos activity will be noted in other areas of the brain known to be necessary for processing social cues. Bulls treated with Depo-Provera maintain sexual interest and have offspring. Depo-Prevera increases the neural activity within the central amygdala and may partially account for their altered aggressive behavior during the rut.

An estimated 90% of reindeer females are mated in a 10- to 21-d interval and give birth in an equally synchronized manner. Reported gestation length in reindeer is highly variable (range, 203 to 240 d), almost twice the reindeer estrous cycle length. Previously, we identified a significant, negative relationship between gestation length and conception date in a small group of reindeer. In the current study, the negative relationship was investigated in a switchback design, where reindeer were divided into two groups synchronized for early and late mating over a 2-yr trial. Regression analysis of 11 paired observations produced a negative (P<0.001) association between gestation length and conception date (slope=−0.31). Dam weight at breeding and prior to parturition, calf birth weight, and calf sex were not significant variables in the regression. Regression analysis of a larger data set from two University of Alaska Fairbanks reindeer herds, where conception date (verified by systemic progesterone) and gestation length were recorded (historical data set), supported previous conclusions (n=70; slope=−0.37; P<0.001). Although the calf sex ratio did not differ with gestation length, there was a positive relationship (r2=0.19; P=0.014) between male birth weight and gestation length in the larger data set. The negative relationship between conception date and gestation length enhanced calving synchrony, though the limits of gestation plasticity and underlying mechanisms are not clear. The potential role of photoperiod on early embryonic development is discussed.

Bovine CIDR were used for 11 d to synchronize estrus in 18 farmed muskoxen. Nine muskoxen went into harem for 1 wk (BRED) following CIDR removal and nine animals served as treated, non-mated controls (CTL). Progestin treatment prevented short estrous cycles (100% BRED; 88.9% CTL). The interval from CIDR removal to CL formation was 7.2 d (BRED) and 11.6 d (CTL) and fertility rate at the estrus following treatment was 88.9% (BRED). The use of bovine CIDR to synchronize estrus reduced the harem period to 1 wk and resulted in synchronized calving (7 d) following a mean gestation of 246 d. Key words: CIDR, Estrous synchronization, male effect, muskoxen, Ovibos moschatus

Reindeer are an important livestock species in Alaska, although the level of reproductive management in reindeer herds is currently low. None-the-less, rapid genetic improvements can be achieved through selective application of AI. Current efforts to develop AI in reindeer require identification of desirable traits worthy of selection (e.g., rapid calf growth rate). Objectives of this trial looked at the relationship between dam BW at parturition, calf birth weight and growth rate (measured as ADG) and the associations between reindeer milk components and calf ADG. Twelve reindeer cows with calves were grouped according to whether the calves were born early (EC; n=6) or late (LC; n=6) in the calving season. Calving dates ranged from April 6 to 11 for EC and from April 23 to 30 for LC. Milk samples were collected from cows at 10, 40 and 70 days-in-milk (DIM). Calf BW was recorded coincident with milk sampling and ADG was calculated for each period and for the entire 70d study. Milk samples were sent to Rocky Mountain DHIA (Logan, UT) for component analysis. Milk energy density was estimated and correlation coefficients were calculated for ADG with dam BW at parturition, milk components and milk energy. Calf ADG did not differ between groups and averaged 0.66 ± 0.06 kg (range 0.53 to 0.89 kg). Dam BW correlated with ADG at 70 DIM (r=.57; p=.04). In the EC group, ADG was correlated with milk energy (r=.92; p=0.01), %fat (r=.91; p=0.01) and %total milk solids (r=.93; p=0.001) at 10 DIM. Milk components appear to be more important to EC calves than LC calves in early life. Reindeer calves are precocious and nibble grass soon after birth. From this study, it appears that milk components play a larger role in early born calves when snow cover prevents access to grass.

Seasonal endocrine changes in 5 non-bred and 10 pregnant Alaskan reindeer have been documented. Blood samples were collected from early September until early May, spanning the breeding season, gestation, or the anovulatory period. Plasma was analyzed by RIA for progesterone (P4), estradiol-17beta, estrone, and estrone sulfate. Elevated P4 in 80% of the reindeer at the onset of the study indicated that ovarian activity had been initiated. The median date for the onset of the first recorded full-length ovulatory cycle was September 23. In nonbred reindeer, the mean ovulatory cycle length from September to May was 24 +/- 1 d (range 18 to 29 d). Nonbred females continued to cycle throughout the winter, displaying 6 to 8 ovulatory cycles after the beginning of blood sampling. Cycle length (mean 22 to 24 d) did not vary between individuals (P = 0.170) or over the course of the winter (P = 0.244). In early April, ovulatory cycles ceased with normal demise of the corpus luteum in 2 females, whereas the remaining 3 females formed apparently persistent corpora lutea. Natural service breeding occurred between September 10 and October 2, and P4 profiles indicated that all breeding females conceived to the first mating. Concentrations of P4 rose steadily after conception and remained elevated throughout gestation, with mean concentrations not varying significantly (P = 0.104) from 4 to 28 wk of gestation. Estrogens all followed patterns similar to each other, remaining at baseline concentrations until approximately 24 wk of gestation and rising coincidently as P4 declined just before parturition. There was a continual overlap throughout the winter in peak P4 concentrations observed in cycling and pregnant reindeer. Calving occurred between April 8 and May 2, resulting in a mean gestation length of 211 +/- 2.2 d (range 198 to 221 d). Information from this study can be used by Alaskan reindeer producers to improve management and profitability of reindeer production.

Muskox farming is an emerging industry in Alaska. For such an endeavor to succeed, it is necessary to develop fundamental husbandry tools specifically for this species. This study examined the role of the bull in the onset of seasonal ovarian activity and the synchrony of estrus and tested the efficacy of a radiotelemetric estrus detection system. Twenty parous female muskoxen, ranging in age from 4 to 15 yr, were divided into three groups (balanced by age and body weight). Groups received early bull exposure (EBE; August 1, n = 7), late bull exposure (LBE; August 22, n = 6), or remained physically isolated from a bull (NBE; n = 7). Just before bull introduction, radiotelemetry transmitters were attached to the rump of muskox cows in the EBE and LBE groups. Plasma samples were collected from all the cows twice weekly and analyzed for progesterone (P4). The mean date of onset of seasonal ovarian activity was earlier in the EBE and LBE groups than in the NBE group (P < 0.001) and was earlier in EBE (P < 0.021) than in LBE. All EBE and LBE cows conceived to the first fertile cycle, giving 100% fertility. The time interval for the onset of ovarian activity between the first and last cow within each group was 7 d in the EBE group, 12 d in the LBE group, and 24 d in the NBE group. Radiotelemetry correctly identified estrus 95% of the time (18 of 19 instances). Mean length of estrus for each cow, calculated from the time of first mount until last mount, was 771 +/- 98 min (range = 4 to 1,508 min). Mean number of mounts during estrus was 8.89 +/- 1.29 (range = 3 to 25 mounts). Length of estrus at the first short cycle did not differ from the length of the first fertile estrus between, or within, individuals. Estrous behavior was initiated more frequently during the night (2300 to 0500; P < 0.05). Bull introduction can be used as a simple, low-cost management tool for synchronizing estrus in farmed muskoxen. Radiotelemetry was an effective tool for identifying breeding behavior in this species.

Reindeer are seasonally polyestrus, short day breeders, with estrous cycles of approximately 20 days in length. The objective of this study was to investigate the effects of reindeer bull exposure on the onset of the reindeer cow breeding season and to investigate whether cows with calving experience responded differently than cows with no previous reproductive experience. During year 1, blood samples were collected weekly beginning 14 July and continuing until 15 September (n = 8) or 30 September (n = 8) in order to determine the onset of the breeding season, based on ovarian function, with no bull present. Plasma was stored frozen for later assay of progesterone (P(4)) following the conclusion of sample collection. Eight randomly selected cows were allowed to breed with a bull during year 1. The mean onset of ovarian activity in the first year was 15 September (range: 8-29 September). The bull was removed from cows more than 2 months prior to the start of the experimental period during year 2 and housed at a separate facility approximately 2 km distant. Blood samples were collected during year 2 from 14 cows 3x weekly beginning on 11 August (day 1) and continuing until 29 September (day 50) and plasma was stored frozen for later assay of P(4) following the conclusion of sample collection. On day 6, cows were divided into two groups such that group 1 (early bull exposure; EBE), consisted of four cows that had calved the previous spring and four cows that had no reproductive experience (n = 8). Group 2 (late bull exposure; LBE), consisted of three cows that had calved the previous spring and three cows that had no reproductive experience (n = 6). EBE experienced bull introduction on day 13, 23 days earlier than the average onset of ovarian activity during year 1. LBE experienced bull introduction on day 46, 10 days later than the average onset of ovarian activity during year 1. Progesterone concentrations were analyzed by ANOVA procedures for repeated measures. Bull presence was not requisite for the onset of ovarian activity during either year. Previous reproductive status had no effect on the onset of ovarian activity within EBE (P = 0.61) or within LBE (P = 0.92). Time of bull introduction had a significant effect on the onset of ovarian activity when EBE was compared to LBE (P<0.001). The first sustained increase in mean P(4) concentration above 1 ng/ml occurred on day 25 in EBE reindeer and day 41 in LBE reindeer. EBE reindeer initiated ovarian activity 12 days after bull introduction while LBE reindeer initiated ovarian activity 5 days before bull introduction. All cows penned with the bull conceived during both breeding seasons, demonstrated by production of live calves during the subsequent spring. Cows bred during year 1 all calved within a 9 day-period. During year 2, EBE displayed a more synchronous calving period compared to LBE (P<0.05). Results indicate that bull management affects the onset of the breeding season in reindeer cows, regardless of previous reproductive experience.

In response to increasing commercial interest and the high market value of qiviut (the downy underwool of the muskox), we have employed standards and measurements used in the wool and cashmere industries to describe qiviut fiber characteristics. Fleece samples (qiviut with guard hair) were shaved from the midshoulder of 299 wild muskox hides of known sex and age (1, 2, 3, and 4+ yr) during the Banks Island, Canada, muskox harvest in November 1997. Samples were analyzed for fiber diameter distribution of raw fiber and qiviut, scoured and qiviut yields, and lengths of guard hair and qiviut fiber. We found a sex x age interaction for average fiber diameter (AFD) in raw fiber (P= 0.002) and qiviut (P < 0.001) only. Adult males had significantly coarser AFD than females (21.5 microm, males vs 20.1 microm, females and 18.2 microm, males vs 17.5 microm, females) for raw fiber and qiviut, respectively. Qiviut AFD from yearlings was 1.7 microm finer than the AFD of adult qiviut. Fiber diameter distribution (SD) decreased with age in the raw sample (P < 0.003) and qiviut (P < 0.001) and qiviut SD was greater (P < 0.001) in males than in females. Qiviut theoretical yield (% mass of fibers < or = 30 microm) increased (P < 0.001) with age, and females had higher theoretical yields than males (P < 0.001). Scoured yield did not vary between sexes in any age class and averaged 93.3%. Qiviut staple length did not differ with either age or sex. In summary, differences between the sexes were small up to the 3rd yr, and these differences were not likely to be of commercial importance. However, considering that AFD is a primary commercial criterion of value, AFD changes from 16.5 microm in yearlings to 18.2 microm in adults and from 17.5 microm in adult females to 18.2 microm in adult males would be expected to result in significant differences in commercial value.

We studied scent marking by adult male caribou (Rangifer tarandus) during rut in September 1998 at the Large Animal Research Station in Fairbanks, Alaska, USA. We used an experimental approach involving two captive groups of two males each to test for effects of social status, tree size, texture, and scent on rubbing behavior by caribou. Dominant males did not rub more often or for a longer duration than subordinates. Caribou rubbed trees with smaller diameters more often than large-diameter trees. Males preferred trees with bark for rubbing to those trees with their bark removed prior to the experiment. Caribou exhibited no preference for posts with pine-oil applied compared with posts without that aromatic scent. We hypothesize that rubbing of trees by male caribou is related to synchronization or priming of estrus in females, but more research is needed to test that potential function of scent marking.