X-linked genes of the H-Y antigen system in the wood lemming (Myopus schisticolor)

Summary H-Y antigen was investigated in 18 specimens representing six different sex chromosome constitutions of the wood lemming (Myopus schisticolor). The control range of H-Y antigen was defined by the sex difference between normal XX females (H-Y negativeper definitionem) and normal XY males (H-Y positive, full titer). H-Y antigen titers of the X^*Y and X^*0 females were in the male control range, while in the X^*X and X0 females the titers were intermediary. Data were obtained with two different H-Y antigen assays: the Raji cell cytotoxicity test and the peroxidase-antiperoxidase (PAP) method. Fibroblasts, gonadal cells, and spleen cells were checked. Presence of full titers of H-Y antigen in the absence of testis differentiation is readily explained by the assumption of a deficiency of the gonadspecific receptor of H-Y antigen. Since sex reversal is inherited as an X-linked trait, genes for this receptor are most likely X-linked. The implications of our findings are discussed in connection with earlier findings concerning H-Y antigen in XY gonadal dysgenesis in man and the X0 situation in man and mouse.     

Sex-chromosome aberrations in wood lemmings (Myopus schisticolor)

The wood lemming displays certain peculiar features: (1) The sex ratio shows a prevalence of females (FRANK, 1966; KALELA and OKSALA, 1966), and some females produce only female offspring (KALELA and OKSALA, 1966). (2) In a considerable proportion (in the present material, slightly less than half) of the females, an XY chromosome complement is found in the somatic tissues, but the Y is absent in the germ line of those studied (Fredga et al., 1976). Therefore, (3) a mechanism of double nondisjunction in early fetal life of XY females has to be postulated, which replaces the Y in the germ line by duplication of the X. It is assumed (4) that the X of XY females bears a sex-reversal factor that affects the male determining action of the Y (Fredga et al., 1977). There is (5) a strong presumption that in most cases the XY females are those that produce daughters only, but (6) a few exceptions may occur (FRANK, unpublished observations), suggesting that the regulation according to assumption 3 (perhaps also to 4) is incomplete in XY females. In the present report, four females are described with a 31,XO karyotype, two females with 33,XYY or 32,XY/33,XYY, respectively, two males with a 33,XXY, and one male with a 32,XX/33,XXY karyotype, as observed in a consecutive series of 502 wood lemmings. The incidence of sex-chromosome anomalies in liveborn and adult animals was 2.3%; the overall incidence, including embryos, was 1.79%. Neither the somatic XO constitution nor the existence of an extra Y in females precludes fertility. However, the XXY condition in the male results in sterility. There is certain evidence that an instability of the proposed mechanism for double mitotic nondisjunction of the sex chromosomes in oogonia accounts for the high rate of sex-chromosome aberrations in wood lemmings, at least when the mother is XY.     

Home range size and activity of the wood lemming, Myopus schisticolor

We present the results from a radiotelemetric study on space use and activity in a natural population of the wood lemming Myopus schisticolor. Male home ranges were larger than females, and the ratio female home range to male home range was smaller than expected compared to other small rodents. Males moved distances 4-12 times longer than females. We argue that this difference in mobility gives a higher probability of capturing males than females in snap trap studies. If there is a sex biased trappability, this might explain the increasing female biased sex ratio during the snow free season (1:1 in spring, 3:1 [female:male| in autumn), as a higher mortality of males during the reproductive season.     

Metabolism,thermogenesis and daily rhythm of body temperature in the wood lemming,Myopus schisticolor

Wood lemmings (Myopus schisticolor) were captured during their autumnal migration in September and October. The animals were maintained at 12°C and under 12L:12D photoperiod. Basal metabolic rate and thermogenic capacity of the wood lemming were studied. Basal metabolic rate was 3.54 ml O₂·g^-1·h^-1, which is 215–238% of the expected value. The high basal metabolic rate seems to be typical of rodents living in high latitudes. The body temperature of the wood lemming was high (38.0–38.8°C), and did not fluctuate much during the 24-h recording. The high basal metabolic rate and the high body temperature are discussed with regard to behavioural adaptation to a low-quality winter diet. Thermogenic capacity, thermal insulation and non-shivering thermogenesis of the wood lemming displayed higher values than expected: 53.0 mW·g^-1, 0.53 mW·g^-1·C^-1 and 53.2 mW·g^-1, respectively. Brown adipose tissue showed typical thermogenic properties, although its respiratory property was fairly low, but mitochondrial protein content was high compared to other small mammals. The 24-h recording of body temperature and motor activity did not reveal whether the wood lemming is a nocturnal animal. Possibly, the expression of a circadian rhythm was masked by peculiar feeding behaviour. It is concluded that the wood lemming is well adapted to living in cold-temperature climates.Abbreviations BAT brown adipose tissue; bm, body mass - BMR basal metabolic rate - C conductance - Cox cytochrome-c-oxidase - HP heat production - HP_max maximum heat production - M metabolism - NA noradrenaline - NST non-shivering thermogenesis - NST_max maximum non-shivering thermogenesis - RMR resting metabolic rate - RQ respiratory quotient - T _a anibient temperature - T _b body temperature - T _lc lower critical temperature - UCP uncoupling protein - vO₂ oxygen consumption - vO_{2 max} maximum oxygen consumption     

Low mitochondrial DNA variation and recent colonization of Scandinavia by the wood lemming Myopus schisticolor

The population genetic structure of wood lemmings ( Myopus schisticolor ) from Scandinavia, Finland and western Siberia was examined by restriction fragment length polymorphism of mitochondrial DNA (mtDNA) in 45 individuals from six localities. The 12 observed mtDNA haplotypes demonstrated a distinct phylogeographic pattern, suggesting that the postglacial colonization of Scandinavia by wood lemmings occurred from north-east. However, a very low level of haplotype and nucleotide diversity and a lack of geographical structure were found within Scandinavia. The limited mtDNA diversity in the Scandinavian populations probably reflects recent divergence in situ after colonization by a limited number of founders. Allozyme data support this scenario.     

On the evolutionary stability of the female-biased sex ratio in the wood lemming (Myopus schisticolor): the effect of inbreeding.

The evolutionary stability of the female-biased sex ratio observed in the wood lemming (Myopus schisticolor) is discussed. The hypothesis analysed is that the skewed sex ratio is maintained as a result of partial and/or recurrent inbreeding. Fredga et al. (1976, 1977) have suggested that an X-linked mutant gene, X, affects the male-determining action of the Y chromosome, thus converting some XY individuals into females. By a mechanism of selective non-disjunction in the foetal ovary only X-carrying eggs are produced. In particular the stability of that genetic mechanism (or the X chromosome) is analysed by considering the introduction of a "suppressing" sex-linked mutant gene Y. Several deterministic simulation models assuming father-daughter and/or brother-sister matings have been developed and analysed. It is concluded that in the case of extremely strong inbreeding, the hypothesised genetic mechanism may, as a result, be evolutionarily stable. Interpreting field observations on microtine rodents in general it is concluded that only a few species are likely to experience such extreme cases of inbreeding. The wood lemming and the related collared lemming (Dicrostonyx troquatus), another case which seems to have XY-females, are likely to exhibit sufficiently strong inbreeding.     

Relationship between nucleoli and sex chromosomes during meiosis of the male wood lemming Myopus schisticolor: a fine-structure study

Electron microscopy of ultrathin serial sections has been used to study the origin and fate of a mass of fibrillar material (FM) during spermatogenesis in the wood lemming Myopus schisticolor. In the course of early pachytene, one of the two nucleoli completely disappears. The remaining nucleolus loses its granular portion and acquires a "round body" encased by the fibrillar moiety, and the restructuring is accompanied by the appearance of FM in the close vicinity of this nucleolus. During diakinesis, the FM increases in volume and density and selectively infiltrates the chromatin of the XY pair. The intermingling of sex chromosomes and FM is at its maximum in metaphase I, giving the XY chromatin a patchy appearance. The FM separates along with the chromatin during the ensuing anaphase I and is shed from the chromosomes during early telophase I. By the time the nuclear envelope is reconstituted, the FM is completely separated from the chromatin. It disintegrates in the spermatids. The FM could not be stained using the Ag-NOR technique. In the wood lemming, X and Y chromosomes show an end-to-end association without a detectable synaptonemal complex. The FM may contribute to the attachment of the two sex chromosomes to each other. Thus, the FM is considered to be a substitute for a chiasma, which normally guarantees proper segregation in anaphase I.     

A new cestode species Paranoplocephala gubanovi sp.n. (Cyclophyllidea: Anoplocephalidae) from the wood lemming Myopus schisticolor of eastern Siberia

A new species of anoplocephalid cestode, Paranoplocephala gubanovi sp. n. (Cyclophyllidea, Anoplocephalidae), from wood lemmings of Eastern Siberia (Myopus schisticolor) is described. The new species differs from other known species of Paranoplocephala associated with Holarctic lemmings by having unique combination of characters as follows: a few-segmented strobila, superficial suckers sticking out of the scolex, ovary covering practically the whole middle part of the segment, relatively little number of testicles situated in the aporal part of the segment, cirrus bursa crossing the poral excretory vessels, and subspherical spermatheca situated in the middle part of the segment. Comparison of P. gubanovi sp. n. and several closest species, P. fellmani Haukisaimi et Henttonen, 2001, P. serrata Haukisaimi et Henttonen, 2000, and P. arctica (Rausch, 1952) has been carried out. Features distinguishing the new species from Aprostatandrya macrocephala and A. microti have also been studied.     

Comparative phylogeography and demographic history of the wood lemming (Myopus schisticolor): implications for late Quaternary history of the taiga species in Eurasia

The association between demographic history, genealogy and geographical distribution of mitochondrial DNA cytochrome b haplotypes was studied in the wood lemming (Myopus schisticolor), a species that is closely associated with the boreal forest of the Eurasian taiga zone from Scandinavia to the Pacific coast. Except for a major phylogeographic discontinuity (0.9% nucleotide divergence) in southeastern Siberia, only shallow regional genetic structure was detected across northern Eurasia. Genetic signs of demographic expansions imply that successive range contractions and expansions on different spatial scales represented the primary historical events that shaped geographical patterns of genetic variation. Comparison of phylogeographic structure across a taxonomically diverse array of other species that are ecologically associated with the taiga forest revealed similar patterns and identified two general aspects. First, the major south-north phylogeographic discontinuity observed in five out of six species studied in southeastern Siberia and the Far East implies vicariant separation in two different refugial areas. The limited distribution range of the southeastern lineages provides no evidence of the importance of the putative southeastern refugial area for postglacial colonization of northern Eurasia by boreal forest species. Second, the lack of phylogeographic structure associated with significant reciprocal monophyly and genetic signatures of demographic expansion in all nine boreal forest animal species studied to date across most of northern Eurasia imply contraction of each species to a single refugial area during the late Pleistocene followed by range expansion on a continental scale. Similar phylogeographic patterns observed in this taxonomically diverse set of organisms with different life histories and dispersal potentials reflect the historical dynamics of their shared environment, the taiga forest in northern Eurasia.     

Gonadal development and birth weight in X*X and X*Y females of the wood lemming, Myopus schisticolor

A quantitative histological analysis of ovaries from 8- to 10-day-old wood lemmings revealed significant differences between females with X*Y and X*X sex chromosome constitutions. The ovarian volume of X*Y females was on average 57% of X*X, and the number of oocytes was less than half in X*Y compared to X*X. However, the frequency of growing oocytes in relation to the total number was 6.5% for X*Y compared to 3.0% for X*X. Oogenesis in X*Y wood lemmings resembles in many respects that of mice heterozygous for certain translocations and with tertiary trisomy (Ts31H), and those with X0 monosomy. The fertility in X*Y wood lemmings is not reduced. On the contrary, X*Y females have a higher reproductive fitness than X*X and XX. This is discussed in relation to the present findings. The body weight at birth was 8% higher in X*Y than in X*X.     

Long-term stability of allozyme frequencies in a wood lemming, Myopus schisticolor, population with a biased sex ratio and density fluctuations

Wood lemming (Myopus schisticolor) populations are characterized by female biased sex ratios and cyclic variations in population size. Both of these characteristics are assumed to reduce genetic variation and thus affect the evolutionary adaptation of the species. We addressed these questions by studying the genetic structure of a wood lemming population from eastern Finland by isozyme markers during a 21-year period, which corresponds to 40-50 generations. Contingency tests showed that genotypic proportions conformed to Hardy-Weinberg equilibrium in each of the four sampling years. Among the temporal replicates, allele frequencies differed most by 0.14 and were not significant. Genetic variation was also stable and fairly high with a mean observed heterozygosity of H = 0.057. Variability in the Heinavesi population was higher than previously reported in wood lemming. The difference was mainly caused by variation at a phosphoglucomutase locus that was monomorphic in earlier studies. Significant linkage disequilibrium was observed in three of the comparisons but the disequilibrium did not appear consistently in all years. This pattern was also evidenced by the variance components, which indicated that selection favoured for specific allele pairs only in few subsamples.     

Correlation between sexual phenotype and X-chromosome inactivation pattern in the X*XY wood lemming

Replication patterns of the X chromosomes were studied in X*XY wood lemmings with male and female phenotypes. The wild-type X was late replicating (ie, inactivated) in all cells of the X*XY female, whereas the mutated X* was late replicating in all cells of the X*XY male. These findings are compared with those obtained in sex-reversed (Sxr) mice.     

Vocal communication in two lemming species (Lagurus lagurus and Eolagurus luteus)

The vocal repertoire of steppe (Lagurus lagurus Pallas, 1773) and yellow steppe (Eolagurus luteus Eversmann, 1840) lemmings consists of only one type of signal, namely, a squeak. The signal does, however, have considerable variability. The DFA standard procedure shows the significant differences in the parameters of squeaks between species and significant differences in squeak parameters between the sexes in steppe lemming but not in the yellow steppe lemming. The fundamental frequency of the smaller steppe lemming’s squeak is higher than that of the larger yellow steppe lemming, although the peak frequencies have comparable values. In addition, the steppe lemming displays a pronounced sexual dimorphism in vocalization parameters; the frequency characteristics of the signals of the smaller females have lower values, which are contrary to scaling laws, stating that smaller individuals tend to have higher fundamental and peak frequencies. The age-related variability in the signals of yellow steppe lemmings was also described. The fundamental frequency of their squeaks is higher than that of the adults, but the peak frequency of signals in young and adults is not distinguished, and its value is higher than theoretically calculated. Such deviation in signal frequency parameters from the optimum may be result of directed selection, indicating the high value of such signals in the acoustic communication of lemmings.     

Variability and identification power of 60 X-chromosome in two native Siberian populations

Genetic diversity of 60 X-chromosome single nucleotide polymorphisms (XSNPid panel) in populations of Siberian Tatars and Tuvinians is described. A close spectrum of allele frequencies and a low level of their genetic differentiation (G_st = 0.021) is revealed. High discriminating power of the XSNPid panel in populations under study is demonstrated. The random matching probability (MP) of multilocus genotypes in males is 1.12 × 10^–18 in Siberian Tatars and 7.77 × 10^–16 in Tuvans. In females, MP is several orders of magnitude lower: 1.51 × 10^–25 in Siberian Tatars and 1.83 × 10^–23 in Tuvinians.     

Cytological identification of cell types in the testis of Esox lucius and E. niger

Summary Testes of Esox lucius and Esox niger were investigated histologically, cytochemically, and ultrastructurally in reproductive fish. Intralobular Sertoli cells possessed numerous lipid droplets in Esox lucius, but not in Esox niger. In both species, interlobular cell types included myoid cells and lipid-negative Leydig cells within the extravascular space. Evidence is presented for a contractile network of myoid cells within the testes of these teleosts. The presence of Leydig cells and myoid boundary cells in the testis of Esox lucius refutes the reported homology between lobule boundary cells and Leydig cells in this species.     

Morphometric variation in the Nearctic collared lemming (Dicrostonyx)

A. H. Macpherson suggested that much of the current geographic diversity in Canadian Arctic mammals resulted from isolation in refugia during the Wisconsin glacial stage. This study evaluates the refugium hypothesis, insofar as it applies to Nearctic Dicrostonyx , by means of a statistical analysis of geographic variation in 13 skull characters. Overall, geographic variation among samples is not significant, although D. hudsonius and D. unalascensis are geographically and morphologically distinct. Some variation in skull shape is correlated with winter temperature. Partitioning tests on other measures of shape variation indicate some discontinuities consistent with the refugial hypothesis. Discrete samples reflect possible refugial populations in northern North America, Eastern Beringia and two southern periglacial refugia, one in eastern North America and at least one in western North America.     

Human lymphokine-activated killer (LAK) cells: identification of two types of effector cells

We analyzed the antigenic phenotype of lymphokine-activated killer (LAK) effector cells. Human blood lymphocytes were cultured for 3 days with 100 U/ml recombinant interleukin 2 (rIL 2), subpopulations isolated with monoclonal antibodies and a fluorescence-activated cell sorter (FACS) and assayed for cytotoxic activity against 51chromium labeled noncultured melanoma tumor cells. Initial experiments compared the LAK effector function of CD5+ T lymphocytes vs CD5- cells (predominantly CD16+ NK cells). The mean percent specific release at a 10:1 effector:target (E:T) ratio was 25% +/- 16 for CD5- cells, 10% +/- 6 for CD5+ cells, and 22% +/- 9 for unsorted cells. In contrast, when lymphocyte subpopulations were isolated before rIL 2 culture (LAK precursors), CD5- cells but not CD5+ cells developed LAK activity (28% +/- 12 vs 1% +/- 1, mean percent specific release, 10:1 E:T ratio), confirming our previous results showing that only CD16+ cells were LAK precursors. The discrepancy between LAK effector and precursor phenotypes suggested that LAK precursors acquired CD5 determinants during rIL 2 culture; however, double label immunofluorescence of rIL 2 cultured CD16+ cells showed that this was not the case. The data suggested that in the presence of other cell types, some T lymphocytes may develop LAK activity, but purified blood T lymphocytes do not develop LAK function when cultured with rIL 2 alone. We also analyzed LAK effector function in lymphocyte subpopulations defined by CD4 and CD8 antigens. The data showed that lymphocytes with a low density expression of CD8 and no expression of CD4 were enriched for LAK effector cells, whereas CD4+ and CD8- had less activity than unsorted cells. Lymphocytes with a high density expression of CD8 had activity similar to unsorted cells. We also assessed the contribution of Leu-7 (HNK-1) granular lymphocytes to LAK effector function. After culture with IL 2, lymphocytes were depleted of Leu-7+ cells by antibody and complement treatment and then were sorted into CD5+ and CD5- fractions. The cytotoxic activity of Leu-7-CD5+ cells was a mean 5% +/- 5 vs a mean 14% +/- 8 for the total CD5+ population (20:1 E:T ratio). The activity of Leu-7- CD5- was slightly less than the total CD5- fraction (21% +/- 9 vs 28% +/- 14, 10:1 E:T ratio). In conclusion, LAK effector function was highest in non-T cell (CD5- CD16+) populations and some activity was also present in T cell populations (CD5+ and predominantly Leu-7+).