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To accommodate a migratory life history, migrants express a greater number of physiological and behavioral stages per annum than residents and are thus considered to have higher finite state diversity (FSD). To investigate the physiological mechanisms and constraints associated with migration, direct comparison of two subspecies of white-crowned sparrow – migrant, Zonotrichia leucophrys gambelii, and resident, Z. l. nuttalli – were made under common garden conditions of photoperiod and housing, as birds progressed from winter through the vernal life history stages. We tested the hypothesis that migrants (higher FSD) respond differently than residents (lower FSD) to the initial predictive cue, photoperiod, to initiate and integrate the progression of vernal stages of prenuptial molt, migration and development of breeding. If differences in vernal phenology were noted, then the basis for the distinctions was considered genetic. Results indicate that (1) residents had a lower threshold to vernal photoperiod with elevations of plasma androgen, growth and development of reproductive structures preceding those of migrants; (2) only migrants displayed prenuptial molt, preparations for migration and migratory restlessness; and (3) neither baseline nor stress-induced plasma corticosterone differed across subspecies, suggesting energetic demands of the common garden were insufficient to induce a differential adrenocortical response in either subspecies, highlighting the impact of environmental conditions on corticosterone secretion. Thus, in a common garden experiment, Z. l. gambelii responds differently to the initial predictive cue, photoperiod, to initiate and execute the vernal stages of molt, migration and development of breeding in comparison to the shared stage of breeding with Z. l. nuttalli, confirming a genetic basis for the subspecies differences.
Migration requires specialized traits of behavior and physiology that are rarely, or never, expressed together in resident species. In spring, most migrants express a sequence of events on the wintering grounds that include prenuptial molt, developmental phases for migration leading to hyperphagia, prominent fuel accumulation, increased hematocrit and flight muscle size, metabolic adjustments to support power and endurance for sustained travel, and activation of the hypothalamic–pituitary–gonadal axis (HPG) for breeding, leading to gonadal recrudescence (Ketterson et al., 2015; Krause et al., 2015b; Morton, 2002; Piersma et al., 2005; Price et al., 2010; Ramenofsky and Wingfield, 2007; Salewski et al., 2009). Furthermore, long-distance migrants cover great distances and encounter a wider range of ecological conditions during their annual movements. By contrast, resident populations of migratory species remain within a more restricted area and are typically exposed to a narrower range of ecological conditions including temperature and availability of food (Alerstam et al., 2003; Gauthreaux, 1982; Mewaldt and King, 1977).
One approach to determine the physiological mechanisms and constraints associated with migration is to directly compare migratory and resident congeners. Migratory (Zonotrichia leucophrys gambelii) and resident (Z. l. nuttalli) white-crowned sparrows offer the unique opportunity for identifying distinctions and similarities into the evolutionary history of migration as well as the regulatory control of the physiological and behavioral traits attributed to the migrant. Historically, both subspecies are thought to have arisen from a migratory ancestor and may have been allopatric for at most 50,000 years (Rand, 1948; Winger et al., 2014). Thus, any distinctions observed between the subspecies represent a long history of separation, indicating that genetic differences in responses to environmental cues that regulate vernal phenology have likely evolved (Pulido, 2007). Here we present an experimental study to parse between genetic and environmental contributions to the migratory and resident life histories.
Comparisons of the subspecies were made under common garden conditions in which photoperiod and housing were identical for both. Populations selected for the study reside at the same latitude (38°N) for 7 months of the year once migrants arrive following autumn migration from the breeding grounds. During these months, resident Z. l. nuttalli concentrate along the coastal zone of northern California while the migrant Z. l. gambelii is more prominent in the Central Valley and foothills of the Sierra Nevada Range of California. Despite exposure to identical photoperiod from October through April, two major differences exist: ecological conditions and respective life histories. Zonotrichia l. nuttalli expresses two life history stages – winter and breeding – while the migrant, Z. l. gambelii, experiences four – winter, prenuptial molt, vernal migration and preparations for the breeding stage (Fig. 1). Such distinctions provide a unique pallet for a comparative study of the role of the environment on life history events of the two subspecies.
According to finite state machine theory (Jacobs and Wingfield, 2000; Wingfield, 2008), species with a greater number of life history stages per annum have higher finite stage diversity (FSD) than those with a fewer number of stages (Colwell, 1974; Jacobs and Wingfield, 2000; Wingfield, 2008; Wingfield et al., 1992a). Therefore, migratory Z. l. gambelii are identified as having higher FSD than Z. l. nuttalli (Wingfield, 2008; Wingfield et al., 1999). Given that all life history stages have a developmental, mature capability and termination phase, each require time to complete. One hypothesis is that organisms with greater FSD have less flexibility in timing and duration of each stage as scheduling is tight. However, with fewer stages, Z. l. nuttalli has more time if one stage is delayed in onset as there is adequate opportunity for its full expression (Wingfield, 2008). Given this, flexibility in timing onset of the developmental stages of the vernal events – breeding for Z. l. nuttalli and prenuptial molt, migration and breeding for Z. l. gambelii – would differ.
Accordingly, individuals with distinct life histories will respond differently to environmental cues. For example, Zonototrichia spp., like many north temperate species, are known to be highly photoperiodic (Farner and Lewis, 1971; Lewis, 1975; Moore et al., 1982; Wingfield et al., 1996, 1997). Specifically, both subspecies held on short-day photoperiods require exposure to the increasing day lengths of spring for activation of the HPG to initiate development of migratory and breeding life history stages (Farner, 1950; Lewis, 1975; Mewaldt and King, 1977; Ramenofsky, 2011). It then follows that migrants with higher FSD should respond differently to the initial predictive cue, photoperiod, to regulate expression of the spring stages than residents with lower FSD. This response allows migrants such as Z. l. gambelii to synchronize timing and progression of seasonal events on the wintering grounds with a timely arrival at the distant breeding site. For the resident, Z. l. nuttalli, residing at only one location, responsiveness to initial predictive information is relevant to initiate development of the breeding stage but with much more flexibility as to when actual nesting begins (Davies et al., 2015; Visser et al., 2009; Wingfield and Silverin, 2009). Therefore, it is possible that Z. l. nuttalli, without the complications of a migratory life history stage, would be far less constrained to initiate development of the breeding stage than Z. l. gambelii (Davies et al., 2015; Visser et al., 2009; Wingfield and Silverin, 2009). If so, Z. l. nuttalli should begin development of the breeding stage earlier than Z. l. gambelii when both are held under common photoperiod and housing conditions. For Z. l. gambelii, this results in a compressed progression through the spring stages to include prenuptial molt that precedes development and expression of the migratory traits. Taken together, we propose that the initiation of the spring events for the two subspecies will be distinct, with the goal to investigate the regulatory mechanisms involved in the precise timing and temporal progression of life history stages of the migratory subspecies.
Considering regulatory mechanisms that may be involved, Lofts and Murton (1968) suggested that the manner by which organisms perceive and respond to day length relates to specific patterns of migration and breeding. For example, species and population differences are based on the degree of sensitivity to day length, which influences the onset and termination of breeding. Species with greater sensitivity – lower thresholds – are those that respond to shorter day lengths, i.e. earlier in spring, before the vernal equinox, whereas those with higher thresholds require longer day lengths to induce an HPG response, allowing development of the spring migratory stage before the breeding stage.
Using an FSD framework, we pose the following hypothesis with four predictions to investigate potential distinctions in mechanisms regulating the development and expression of the vernal life history stages of the two subspecies. Migrants with higher FSD respond differently than residents with lower FSD to the initial predictive cue, photoperiod, to initiate and integrate the progression of vernal stages of prenuptial molt, migration and development of breeding. If the differences in vernal phenology are genetic, then in a common garden of environmental conditions, both subspecies will maintain their dissimilarities in timing and temporal progression of vernal events. By contrast, should environmental conditions dominate, then no distinctions across the shared stages will be apparent.
The four predictions are as follows. (1) Gonadal activity: residents will show a lower threshold of gonadal development to increasing spring photoperiods than migrants. Therefore, growth and development of reproductive organs, cloacal gland and testes of residents will be advanced over those of migrants. (2) Body condition: migrants, but not residents, will display molt, fattening, body mass increases, flight muscle hypertrophy and elevated hematocrit in preparation for migration. (3) Steroid hormones: as the life history stages of the subspecies diverge throughout the progression of the vernal events, the timing and magnitude of androgen and corticosterone secretions in residents will exceed those of migrants. (4) Migratory behavior: nocturnal migratory restlessness will be displayed only by migrants.
Field methods
Seed-baited potter traps and Japanese mist nets were used to capture adult male Z. l. gambelii (N=12) on their wintering grounds in Yolo County, California (38°33′N, 121°44′W), and adult male Z. l. nuttalli (N=11) along coastal Sonoma County, California (38°31′N, 123°06′W), between November and December 2012. Body size was assessed by measurements of the relaxed wing chord and tarsus length using calipers (to the nearest 0.1 mm). Sex was determined initially by wing length and confirmed subsequently by unilateral laparotomy.
Laboratory conditions
Birds were brought into captivity and held in quarantine for 1.5 weeks in flight aviaries, after which each subject was placed in individual registration cages (35×40×45 cm, width×length×height) equipped with a photo-detector (Alarm Entry Device, RadioShack Corporation; Forth Worth, TX, USA) positioned 0–10 cm above a central perch and 34 cm above the bottom of the cage. Once a bird crossed the infrared beam emitted from the detector, an electronic signal was transmitted to a Mini Mitter Acquisition System – Vital View (Sun River, OR, USA). Continuous 24-h locomotor activity was recorded in 15 min bins and analyzed in hourly intervals per week for each bird over the entire 5 month study. The photoperiod was adjusted to track natural day lengths of 38.5°N using a Paragon EL72PC digital timer with daily temperatures ranging from 18 to 23°C. Birds were supplied with food ad libitum in the form of commercial bird seed containing millet, cracked corn and sunflower seeds (Mazuri Small Bird Maintenance Diet, 56A6; www.mazuri.com) and Health Blue Grit (Seed Factory, Ceres, CA, USA). Fresh drinking and bath water were provided daily. Nocturnal locomotor activity is not expressed in total darkness (Coverdill et al., 2006; Ramenofsky et al., 2008); therefore, low intensity illumination (Limelite Nightlight, Austin Innovations, TX, USA, <1.0 lux at the source measured by a Greenlee Digital Light Meter (93-172, Rockford, IL, USA), was positioned centrally in the room and visible to all cages.
Captive methods (14 December to 2 May)
Once birds acclimated to individual cages, collection of morphological data commenced on 21 December (noted as week 1) and was repeated approximately twice monthly at the beginning and the middle of each month starting 3 h after sunrise (Table 1). First, body mass was determined to the nearest 0.1 g with a Pesola scale. Fat score was assessed visually by observing presence of lipid in the coelomic cavity and chorico-clavicular fossa with values that ranged from 0 (no observable fat) to 5 (bulging fat deposits) (Ramenofsky and Németh, 2014). Cloacal protuberance (CP) length was measured with calipers to the nearest 0.1 mm. Condition of the flight muscles (pectoralis complex) was determined with a visual score of the muscle profile. Scores were based on four classes ranging from 0 to 3 following the methods of Bairlein (1994). Specifically, a score of 0 presents a sharp protruding edge of the keel to touch and a notable concave muscle mass that is typical of emaciated birds. A score of 1 has a sharp keel edge but the muscle bulges out slightly. A score of 2 has muscle that is convex in shape but the keel edge can be felt between both left and right muscle masses. And with a score of 3, the fibers of the pectoralis rise above the keel that is now embedded in the bulging muscle.
Migratory restlessness
Migratory restlessness, also called Zugunruhe, is the elevated nocturnal locomotor activity expressed in captive birds during the migratory periods that corresponds with the migration of free-living congeners (Berthold and Querner, 1988; Farner et al., 1954; Ramenofsky et al., 2003). Extensive analyses of the 24 h locomotor recordings of both migrant and resident white-crowned sparrows have identified distinguishing features during the spring migratory stage: (1) maximal nocturnal activity exceeds that of the photophase (light phase) (Coverdill et al., 2006; Ramenofsky et al., 2008) and (2) expression of the intense nocturnal activity commences with onset of the scotophase (dark phase). In the present study, we used these criteria to verify that the elevated nocturnal locomotor activity was in fact expression of migratory restlessness in the spring migrants.
Presence of prenuptial molt was determined by scoring three body regions in both subspecies: crown, back (including nape, back and rump) and abdomen (including throat, breast, abdomen and flanks) following Ramenofsky and Németh (2014). Each region was given a score between 0 and 3 depending on the extent of molting feathers: 0, no molt (0%); 1, light molt (1–10% area); 2, moderate molt (11–50% area); and 3, heavy molt (51–100% area). Scores from each region were summed to generate a total molt score for each bi-weekly measurement.
Blood sampling
Blood samples for corticosterone (CORT) and androgens [testosterone and 5α-dihydrotestosterone (DHT)] were collected on five dates approximately once per month starting the week of 15 January (noted as week 4; Table 1). For CORT, collection of the first blood sample was completed within 3 min of entering the room and represented baseline-level circulating glucocorticoid prior to the disturbance. Testosterone and DHT were measured in a second blood sample taken within 10 min of the disturbance. The standard stress restraint protocol was used to measure the second or stress-induced CORT sample that was collected 30 min after the initial disturbance and considered the active response to the capture and handling procedure (Wingfield et al., 1992b). All samples were collected into heparinized micro-hematocrit tubes. Samples were centrifuged for 5 min to separate the red blood cells from plasma according to Fair et al. (2007), and hematocrit was measured for every capillary tube from the baseline samples and then averaged to yield a mean hematocrit value following methods of Krause et al. (2015b). Plasma was subsequently aspirated and stored at −30°C until assayed according to Ramenofsky and Németh (2014).
CORT, DHT and testosterone radioimmunoassays
Thawed samples were treated with a 2000 CPM dose of each 3H DHT and testosterone to determine percent recovery for each sample. Steroids were extracted with ethyl ether, dried under nitrogen, reconstituted in 5% ethyl acetate in iso-octane and added to diatomaceous earth/glycol (50/50 ethylene and propylene glycol) microcolumns for partial purification. DHT was collected first in 10% redistilled ethyl acetate in iso-octane followed by testosterone in 20% ethyl acetate in iso-octane (Wingfield and Farner, 1975). Samples were dried under a stream of N2 and reconstituted in 550 µl PBSG. Procedures for the preparation and extraction of the corticosterone were identical to those described above except that 2000 CPM of 3H CORT was added to each sample for recovery determination and the extracted and reconstituted samples were put directly into the radioimmunoassay (RIA) because the antibody shows no cross-reactivity with other glucocorticoids present in white-crowned sparrow plasma samples (Landys et al., 2004; Wingfield and Farner, 1975). All samples were run in duplicate in multiple RIAs using DHT and testosterone curves that ranged from 3.9 to 1000 pg and CORT from 7.8 to 2000 pg (Landys et al., 2004; Ramenofsky and Németh, 2014). Recoveries for each steroid ranged as follows: DHT, 47–87%; testosterone, 30–70%; and CORT, 67–89%. The sensitivity of each steroid (as determined by the water blanks) and inter-assay variation were 0.33 ng per tube and 10.9% for DHT, 0.24 ng and 15% for testosterone, and 1.04 ng and 5% for CORT. Samples were combined with Ultima Gold scintillation fluid and then counted for 6 min or to within 2% accuracy on a Beckman 6500 Scintillation Counter. Final concentrations of each sample were calculated in units of ng ml−1 of plasma following corrections for plasma volume and recovery counts. Labeled testosterone [1,2,6,7,16,17-3H(N)] (NET553250UC) and DHT (5α Androstan-17β-OL-3-One) [1,2,4,5,6,7-3H(N)] (NET453250UC) were purchased from PerkinElmer (Waltham, MA, USA) and rabbit anti-testosterone-3 (20R-TR018w, Lot 01916) from Fitzgerald Industries International (Acton, MA, USA). 3H CORT was purchased from PerkinElmer (NET 399250UC) and CORT antibody from Esoterix (B3-163, Calabasas Hills, CA, USA).
Gonad inspection by unilateral laparotomy
To confirm gender and determine testis length, unilateral laparotomy was conducted at the conclusion of the study (week 19). Each bird was deeply anesthetized with 2–4% isoflurane in oxygen via a Summit anesthesia vaporizer (Bend, OR, USA). Next, a small incision between the last two ribs provided visual access to the gonad, where length was measured. The incision was sealed with Nexaband Liquid (Abbott Labs, Chicago, IL, USA). Upon recovery, an intramuscular injection of Meloxicam-Metacam (5 µg kg−1, Boehringer Ingelheim, Fremont, CA, USA) was administered at a dose of 0.1 mg ml−1 in sterile normal saline (Vedco Inc., St Joseph, MO, USA) for analgesia. Birds were observed postoperatively over the next 7 days. All procedures and animal handling were done in accordance with the University of California Davis Institutional Animal Care and Use Committee (IACUC protocol 17144) and conducted under the scientific collecting permits issued by the California Department of Fish and Wildlife (11024) and the US Fish and Wildlife Department (MB11826A-5).
Statistical analyses
All analyses were performed either in R version 3.2.4 (R Foundation for Statistical Computing, Vienna, Austria) with the glmmadmb package (Fournier et al., 2012) or SPSS Statistics Software (version 22.0, IBM, Armonk, NY, USA). First, morphological, hematocrit and hormone values were analyzed by fitting linear mixed-effects models (LMM) with sampling week, subspecies and their interactions as main effects and bird ID as a random effect accounting for the repeated sampling design. Distributions of variables were tested for normality with the Shapiro–Wilk test initially or after plotting the residuals versus the predicted values. Records of body mass represent a change from the initial measurement of each bird recorded on the first date of the study from that collected on the subsequent dates. Categorical data – scores of muscle profile – were tested with a generalized linear mixed-effects model for repeated measures using a Wald chi-square statistic. For comparisons of the nocturnal locomotor activity of migrants and residents throughout the study, activity of the first 7 h of the dark or scotophase were averaged by week for each individual prior to analysis. Activity was modeled using a generalized linear mixed-effects model using a negative binomial distribution ('nbinom1' in glmmadmb) with a random effect of individual to account for repeated sampling. The model included fixed effects of subspecies, week and their interaction. To verify that the elevated nocturnal activity recorded in the migrants from experimental week 15 onwards represented migratory restlessness, comparisons of activity measured during the photo and scotophases were drawn both within and across subspecies before and after the appearance of heightened nocturnal restlessness with paired and unpaired parametric and nonparametric tests, Wilcoxon signed-rank or Mann–Whitney U-tests, as required. All post hoc tests comparing the subspecies were conducted with either a Student's t-test or Mann–Whitney U-test with Bonferroni corrections and level of significance across all tests was set at P<0.05.
Molt and body condition
Prenuptial molt was first noted during week 11 but only in migrants, and peaked by week 15 (P<0.05) and then subsided (Table 2, Fig. 2A). The three measures of body condition included changes in body mass, fat score and flight muscle profile. Mass and fat score increased in migrants over residents by week 13 (P<0.05; Tables 2, 3, Fig. 2B–D), but flight muscle score rose only in the migrants by week 17 (P<0.05; Fig. 2D).
Blood components and reproductive structures
Hematocrit increased in both subspecies without distinction between the subspecies (P>0.05; Table 2, Fig. 3A). Elevations of plasma testosterone were first noted in Z. l. nuttalli in week 8, with DHT rising 4 weeks later (Table 2, Fig. 3B,C). Zonotrichia l. gambelii showed no elevation of androgens until the end of the study (week 19), emphasizing the distinct pattern of androgen secretion, with residents showing earlier onset and greater magnitude increase. Changing in parallel with plasma steroids, CP of the residents started increasing by week 11 and continued throughout the remainder of the study, whereas the migrants initiated only a modicum of growth by week 19 (Table 2, Fig. 2E). Testis length assessed by laparotomy at the conclusion of the study (week 19) was greatest in Z. l. nuttalli (t-test: t1,22=−6.59, P<0.001; Table 4). There were no significant correlations of circulating levels of androgen with either morphological measure of CP height or testis length in either subspecies (P>0.14 in all cases). Neither measure of plasma CORT – baseline or stress-induced – changed throughout the study or differed across the subspecies (P>0.05; Fig. 3D,E).
Migratory behavior
Starting in week 1, low-level nocturnal locomotor activity was detected in both subspecies until week 15, when the frequency increased significantly in migrants, reaching a peak in week 17 (P<0.05; Table 4, Fig. 4A). To verify that this activity represented migratory restlessness, we used the criteria established previously for Z. l. gambelii (Agatsuma and Ramenofsky, 2006; Coverdill et al., 2011; Ramenofsky and Németh, 2014). Prior to week 15, the 24 h locomotor activity patterns of both subspecies were similar (Fig. 4B). Pair-wise comparisons of activity recorded during the scoto- and photophases were distinct (Z. l. gambelii, T =−4.1, n=11, P=−0.002; Z. l. nuttalli, T =−8.41, n=10, P=0.000). But from week 15, the nocturnal locomotor activity exceeded that of the photophase in Z. l. gambelii (Z=−2.12, P=0.03, n=12), whereas photophase activity exceeded that of the scotophase in Z. l. nuttalii (Z=−2.9, P=0.003, n=11; Fig. 4C). Furthermore, pair-wise comparisons of mean hourly activity across the last 3 h of the photophase and the first 3 h of the scotophase detected a significant increase at onset for Z. l. gambelii (T =25.6, P=0.002), without any change observed in Z. l. nuttalli (T =−2.77, P=0.11).
Comparisons of migratory and resident subspecies of white-crowned sparrow reveal insights into the physiological mechanisms and constraints associated with migration. With regards to finite state machine theory (Ramenofsky and Wingfield, 2006; Wingfield, 2005), this study focuses on prenuptial molt and developmental phases of migratory and breeding life history stages. Fig. 1, drawn from field studies of Zonotrichia, illustrates progression of the vernal stages during which both subspecies inhabit the same latitude. As straightforward as this figure appears, what has not been tested directly is how the two subspecies prepare for the vernal stages under common garden conditions using the initial predictive cue – photoperiod. Consistent with earlier studies that compared the effects of day length with short-day controls in both subspecies (Blanchard, 1941; Farner, 1974; Lewis, 1975; Mewaldt and King, 1977), our results provide support for the hypothesis that both migrants and residents respond to onset of spring photoperiods, but the timing of the response differs. Development of the breeding stage of Z. l. nuttalli precedes that of the migratory and breeding stages of Z. l. gambelii. By contrast, the local predictive cue, temperature, had minimal effect on photoperiodically induced gonadal growth in Z. l. gambelii, whereas both temperature and precipitation altered the rate of recrudescence in Z. l. nuttalli (Blanchard, 1941; Wingfield et al., 1992a, 1996). This suggests that local (environmental) conditions have a greater impact on the rate of progression of breeding cycles in resident subspecies with lower FSD. However, this resident/migrant distinction may not hold true for an altitudinal migrant such as the western song sparrow (Melospiza melodia morphna), which winters at low elevation (0–10 m) and breeds in the Cascade Mountains of Washington State (500–1220 m) (Perfito et al., 2005). Exposure to low mountain temperatures slowed the rate of gonadal growth in comparison to low elevation temperatures, demonstrating the influence of conditions on the breeding grounds for a short-distance migrant that has the option to retreat until conditions improve on the breeding grounds. This would not be the case for arctic breeders such as Z. l. gambelii.
Prenuptial molt
The first stage to develop in migrants was prenuptial molt, without observable changes in Z. l. nuttalli (Fig. 2A). In juvenile Z. l. gambelii, this molt serves to replace brown and white crown feathers with adult black and white plumage. Birds also replace body feathers to support insulating properties and protection from the environmental wear and tear experienced throughout migration and breeding (Jenni, 1994; King, 1968). Increased vascularization of the feather follicles, protein synthesis, and alterations in concentrations of blood and bone components also accompany feather replacement (Wingfield and Silverin, 2010). Thus birds incur major costs of time and energy but the contributions supporting body condition for ensuing stages are critical. Lack of a complete prenuptial molt in Z. l. nuttalli may be a result of the energy or time required for the breeding stage outstripping the need for a plumage upgrade. Black-crown males hold larger territories, yet brown-crown birds pair and breed, suggesting that age-related morphology may not constrain breeding success (Ralph and Pearson, 1971). Regulation of prenuptial molt is not well understood (Wingfield and Silverin, 2002), but Moore et al. (1982) showed that Z. l. gambelii held on short photoperiods (8 h:16 h light:dark) failed to molt while those maintained on constant photoperiod of 12 h:12 h light:dark did. In the present study, molt was first observed when photoperiod reached 11 h:13 h light:dark (Table 1), which may pinpoint a threshold for molt initiation in Z. l. gambelii (Fig. 2A). Thus, prenuptial molt is important for bolstering body condition in migrants facing long-distance travel but requires time for progression of the stage preceding full preparation for migration.
Body condition
A major component of migratory preparation includes a notable increase in food intake – hyperphagia – that results in increased body mass and fat deposition (Morton, 1967; Ramenofsky and Wingfield, 2007). Fat (lipid) is a primary fuel that migratory species use for long-distance flight (Jenni and Jenni-Eiermann, 1998; McWilliams et al., 2004; Price et al., 2010; Ramenofsky, 1990). As mass and fat score peaked, flight muscle profile notably increased in Z. l. gambelii (Fig. 2D). Such enlargement or hypertrophy is due to increased size of flight muscle fibers and lipid deposition supporting enhanced power and fuel for migration, yet mechanisms controlling its development are largely unknown (Bauchinger and Biebach, 2005; Cornelius et al., 2013; Dietz et al., 2007; Gaunt et al., 1990; Marsh, 1984). Results from a comparative study of resident and migratory dark-eyed juncos (Junco hyemalis) during spring identified differential expression of genes for metabolism and transport of lipids in flight muscle of migrants (Fudickar et al., 2016b). Further evidence for enhanced utilization of lipid by flight muscle in migrants comes from the seasonal upregulation of fatty acid transporters, phospholipid composition, as well as oxidative enzymes in flight muscle of migratory white-throated (Zonotrichia albicollis) and white-crowned sparrows (McFarlan et al., 2009; Price et al., 2010). Taken together, these findings provide support for enhanced body condition (prediction 2) in migrants as birds prepare for spring departure.
Measures of hematocrit provide information about the oxygen-carrying capacity of blood and can be considered an indicator of the responsiveness to metabolic requirements. Previous findings report similar patterns of hematocrit in free-living populations of Z. l. gambelii and Z. l. nuttalli until levels escalated in migrants upon arrival on breeding grounds in May (Krause et al., 2015b). This may be explained by the increased energetic demands associated with migration and preparations for breeding. Yet in captivity, the photoperiodically induced changes in hematocrit generated comparable levels in both subspecies regardless of whether breeding or migratory features were expressed. By contrast, Chilgren and deGraw (1977) found a negative relationship of hematocrit with molt intensity in Z. l. gambelii. It is possible that our blood sampling schedule was insufficient to observe the more discreet changes in hematocrit. Nevertheless, unlike the other measures of body condition, no distinction across the subspecies of hematocrit was identified.
Timing of development and expression of breeding
Previous ecological studies noted the earliest recorded dates for presence of sperm in the seminiferous tubules of the testis and actual matings for Z. l. nuttalli was 4 March (Blanchard, 1941; Mewaldt and King, 1977). However, more recent observations pinpoint mid to late March (Krause et al., 2015b). Both gonadotropins and testosterone are required for spermatogenesis (Ubuka et al., 2014), and our results show that plasma testosterone increases in Z. l. nuttalli by week 8 and DHT increases 4 weeks later (Fig. 3B,C). Using these data as benchmarks and considering that spermatogenesis requires at least a month for full development of spermatids, we calculated back to suggest that activation of the HPG may occur as early as January and possibly even late December, as suggested by Blanchard (1941) and Mewaldt et al. (1968). Thus, the total timing and magnitude of the androgen secretion observed in the resident throughout the study exceeded that of the migrant, and offers support for the androgen profile component of the steroid hormone prediction (prediction 3). It is also possible that Z. l. nuttalli may be less sensitive than Z. l. gambelii to some inhibitory environmental factors affecting the transduction of the photic signal impinging upon thyroid hormones and deiodinase 2. This could alter the hypothalamic–pituitary–thyroid axis shown to regulate photoinduction of breeding in Japanese quail, Coturnix coturnix japonica (Yoshimura et al., 2003). In support, chemical inhibition of thyroid hormones impaired the development of migratory and breeding traits in Z. l. gambelii (Perez et al., 2016).
Reproductive structures
The cloacal protuberance (CP) functions to store and support delivery of sperm during copulation. Hypertrophy of the structure depends upon the seasonal increase of androgen, particularly the steroid action of DHT at the target tissue (Balthazart, 1983), and may serve as a qualitative measure for plasma levels of testosterone. Enlargement of the CP in Z. l. nuttalli preceded that of Z. l. gambelii, providing additional evidence for the earlier developmental phase for breeding in the resident and support for the timing of gonadal activity prediction (prediction 1) (Fig. 2E). However, neither plasma testosterone nor DHT were significantly correlated with CP length, suggesting that development of this reproductive structure may be affected more at a threshold level rather than at discreet values of androgen. At the conclusion of the study, testis length of residents exceeded that of migrants (Table 4). However, such measures collected upon arrival at a northerly site of the breeding range (9.52±0.55 mm, mean±s.d.) exceeded those of the captive migrants (7.9±0.71 mm; t1,17=7.49, P<0.0002), indicating that development had yet to be achieved in the captive migrants. No correlation was found between plasma levels of testosterone or DHT and testis length in either subspecies, suggesting that both testicular recrudescence and spermatogenesis occur long before plasma levels of androgen peak.
Migratory behavior
A key measure of an individual's propensity to migrate in captivity is the expression of nocturnal migratory restlessness in captive birds (Agatsuma and Ramenofsky, 2006). This behavior first appeared in Z. l. gambelii by week 15 and continued to rise over the next 2 weeks (Fig. 4A). Onset of this activity was commensurate with the timing of migration in free-living birds throughout Yolo County (M.R. and Z.N., field observations). Prior to the appearance of migratory restlessness, both subspecies showed elevated activity early and late in the photophase coinciding with bouts of feeding (Agatsuma and Ramenofsky, 2006; Coverdill et al., 2011; Ramenofsky et al., 2008). Such observations highlight the behavioral and physiological similarities of the two subspecies during the winter stage. However, once migrants initiated migratory restlessness, the activity patterns diverged, providing support for the migratory behavior prediction (prediction 4).
Plasma steroids in relation to behavior and physiology
In light of the differential response to spring photoperiods, plasma levels of androgen in Z. l. nuttalli preceded and exceeded those of Z. l. gambelii by 7 weeks. If one considers onset of migratory restlessness as timing of departure (week 15, Fig. 4A), plasma levels of testosterone and DHT remained unchanged from winter, suggesting that androgens do not regulate the switch from diurnal to nocturnal activity in Z. l. gambelii, as proposed for European starlings (Sturnus vulgaris), dark-eyed juncos and gray catbirds (Dumetella carolinensis) (Owen et al., 2014; Subbaraj and Gwinner, 1985; Tonra et al., 2011). Identifying factors that regulate onset of migratory flight are enigmatic, but the neurosteroid 7α-hydroxypregnenolone may hold promise for its role in the diurnal locomotor activity in Japanese quail (Coturnix japonica) (Tsutsui et al., 2008, 2012).
Earlier timing of androgen elevations in Z. l. nuttalli offers support for a genetic basis underlying a lower threshold to photoperiod for the onset of breeding in comparison with Z. l. gambelii (Blanchard, 1941; Mewaldt et al., 1968). Zonotrichia l. nuttalli may well rely on local predictive cues (temperature, food availability) for regulation of the progression through the breeding stage without the complication of the added migratory life history stage. Should conditions degrade during development of the stage, Z. l. nuttalli may slow the rate of progress to onset of breeding given the more relaxed schedule and option for successive broods, neither of which is possible for Z. l. gambelii. Other comparative studies of migratory and resident species have recorded earlier onset of breeding parameters as well as earlier gene expression for reproductive mechanisms among residents, offering additional support for differential regulatory mechanisms relating to FSD (Atwell et al., 2014; Fudickar et al., 2016b; Grieves et al., 2016).
Furthermore, we found no support for the prediction that CORT levels would differ in the two subspecies across the stages. The hypothalamic–pituitary–adrenal (HPA) axis responds to increased energetic demands associated with life history stage and perturbations in the environment (Eikenaar et al., 2015; Landys et al., 2006; Romero and Wingfield, 2015). In free-living birds, residents have slightly higher baseline and stress-induced levels of CORT during the winter months than migrants (Krause et al., 2014). Such differences in CORT in free-living birds may be attributed to differences in either behavior or environmental conditions between the two field sites. Comparing the local habitats for each subspecies – coastal versus the Central Valley – each differs in terms of structure, floral and faunal communities, as well as climate, all of which likely contribute to differential demands and responses of the HPA axis. Furthermore, stress-induced levels are elevated upon arrival in spring and onset of breeding in free-living Z. l. gambelii, whereas Z. l. nuttalli show a more muted response at the outset of their breeding stage (Addis et al., 2011; Krause et al., 2015a, 2016). However, under our common garden conditions without the imposition of natural environmental demands, birds showed similar baseline and stress-induced responses. By contrast, other studies of congeners or partial migratory populations report distinctions in either baseline or stress-induced levels of CORT, suggesting selection, developmental processes or season of sampling as possible causes (Angelier et al., 2011; Eikenaar et al., 2015; Fudickar et al., 2016a). Nevertheless, we found no support for the prediction that CORT levels would differ in the two Zonotrichia subspecies across the 5 months of the study, pointing to a strong influence of environment conditions on adrenal secretion.
Our results confirm that Z. l. gambelii (high FSD) responds differently than Z. l. nuttalli (low FSD) to identical environmental conditions mimicking vernal phenology and support the premise that these differences across the subspecies have a genetic basis. The initial predictive cue, changing photoperiod, initiates and integrates the temporal progression of the spring stages – prenuptial molt, vernal migration and development of breeding – in Z. l. gambelii. In contrast, Z. l. nuttalli showed early gonadal recrudescence and maturation in the absence of competing life history stages (namely, molt and migration). These findings are consistent with the hypothesis that species with greater FSD have less flexibility in timing and duration of each stage given the greater number of stages to be completed within a year compared with species with lower FSD. The primary mechanism involved with the distinction is likely the increased sensitivity of the cue response system of Z. l. nuttalli to a shorter photoperiod following the annual nadir of winter solstice. This results in an earlier onset of the developmental phase of the breeding stage. Local and short-term environmental conditions may have contributed to hasten the earlier onset in Z. l. nuttalli because in a resident population, local conditions are relevant for the timing of breeding. For migrants wintering in locations distant from their breeding grounds, timing of prenuptial molt and migration prior to reproductive development requires photoperiodic information, perhaps the most precise and consistent environmental cue. The vernal increase in day length maintains the pace of the life history stages so that birds arrive on breeding grounds at an appropriate time. The local predictive cues on the wintering grounds during the preparatory phases of the vernal life history stages would be largely irrelevant for timing. However, once on the breeding grounds, local conditions likely predominate to regulate progression of breeding. Mechanisms behind these distinctions are not known but must involve perception, neuroendocrine transduction and response to the threshold levels of photoperiod (Wingfield, 2015). The result of these distinctions set the two subspecies on separate trajectories, with Z. l. nuttalli initiating and progressing through breeding earlier in the year while Z. l. gambelii requires a longer photoperiod to initiate development of molt, migration and gonadal recrudescence. This delay enables accurate scheduling for arrival on breeding grounds coincident with conditions conducive for reproduction.
Given the photoperiodic responsiveness and tight scheduling for a long-distance migrant, delays or advances of phenological conditions in spring on the breeding grounds would result in a mismatch of breeding readiness and optimal environmental conditions. Whether Z. l. gambelii with its high FSD is able to adjust remains uncertain compared with Z. l. nuttalli, with a lower FSD. The stage is now set for experiments manipulating initial predictive and local predictive cues in spring to determine whether resident Z. l. nuttalli integrates more environmental cues than Z. l. gambelii and to identify the neural pathways involved.
We dedicate this work to the inspiring life and work of Professor Peter Marler, whose support to M.R. contributed to the beginning forays into the physiological and behavioral approaches to the studies of white-crowned sparrow migration. We appreciate the ideas and support for the many aspects of this study provided by Roni Chau, Tom Hahn and John Wingfield. Tom Hahn generously conducted the laparotomies at the conclusion of the study.
Competing interests
The authors declare no competing or financial interests.
Author contributions
Conceptualization: M.R., Z.N. Methodology: M.R., A.W.C., Z.N. Software: J.H.P., J.S.K. Validation: M.R. Formal analysis: J.H.P., J.S.K. Investigation: M.R., A.W.C., Z.N. Resources: M.R. Data curation: Z.N. Writing - original draft: M.R. Writing - review and editing: M.R., A.W.C., J.H.P., J.S.K., Z.N. Visualization: M.R., J.H.P., J.S.K. Supervision: M.R., Z.N. Project administration: M.R. Funding acquisition: M.R.
This work was supported by grants from the National Science Foundation to M.R. (IOS-0920791, ARC-1147289).
- Received August 17, 2016.
- Accepted January 25, 2017.
- © 2017. Published by The Company of Biologists Ltd


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subset: commoncrawl


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Needham: St Peter
Our website is: www.7churches.org.uk
Announcement on the Re-Opening of Churches The Government has announced that church buildings may re-open for worship from Saturday 4th July. However, as Covid 19 has not gone away there will be limitations on church opening for some time to come. Whenever one of the 7churches is open strict safety measures will be in place. Online services will be continuing.
On Sunday 5th July the following churches will be open for worship, and will remain open afterwards for Private Prayer until 4pm.
Brockdish (St. Peter & St. Paul's): 9.30am Holy Communion
Harleston (St. John's): 11am Holy Communion
Scole (St. Andrew's): 11am Holy Communion
Safeguarding: The parish of Needham: St Peter is committed to the safeguarding of children, young people and adults. We follow the House of Bishops guidance and policies and have our own Parish Safeguarding Officer(s), PSOs. The Diocese of Norwich's safeguarding pages contain vital links and information including contacts for the Diocesan Safeguarding Advisor (DSA) who advise our PSOs. If you are concerned that a child or adult has been harmed or may be at risk of harm please contact the DSA. If you have immediate concerns about the safety of someone, please contact the police and your local authority Children or Adults Services.


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According to KDSK-TV, Martin L. Evanick, the 31-year-old drummer and vocalist for the Illinois death metal band THE CORE OF YOUR NIGHTMARES, is facing federal child pornography charges.
Evanick, who performs under the pseudonym "Vlad," was indicted last week and is being held without bail.
According to Southern District of Illinois prosecutors, Evanick had sex with a minor in April and took close-up photos of the minor's genitalia.
If convicted, Evanick faces between 15 and 30 years in prison, a $250,000 fine and term of supervised release between five years and life.
The other members of THE CORE OF YOUR NIGHTMARES posted a statement on Facebook distancing themselves from Evanick's behavior. "We had no involvement or knowledge about these acts, and we in no way condone in such actions," they wrote. "What happened to him is unfortunate, but he brought it on himself."
They added in a separate post, "We DO NOT condone in any of the actions Vlad partook and we are appalled by his behavior. [He] never exhibited any kind of behavior that would make us suspect he would do what he did."
Evanick's THE CORE OF YOUR NIGHTMARES bandmates have vowed to continue making music without him under another name.


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Pretty much for as long as I can remember my youngest son has had struggles when it comes to eating and going potty. When he was a baby he would constantly throw up after he ate. I can't even tell you how many outfits the two of us went through in a day. His doctor at the time didn't really have an answer as to why this was happening so we just kept pushing through.
Fast forward to around 6 months of age, time to try out solids. We decided to start with avocado mixed with breastmilk. He wasn't terribly interested but wasn't opposed either. He gagged of course but that's normal for babies who are just starting out so we did t think anything of it. From that point on every food we tried he would gag. Eventually we realized this wasn't your normal baby learning to eat gagging. We talked to the doctor about our concerns, we did the wait and see for a bit first because maybe he just wasn't quite ready but it just didn't get better. At one years old he still wasn't eating. When he did try he would gag or choke, it was a stressful time to say the least.
We ended up with a referral to the children's hospital. At this point not only was he not eating but he was going a week or more without pooping. When he could finally go it was a thing over nightmares. It is crippling to see your child in that kind of pain. So for the next year it was test after test and biopsy after biopsy. We got to the point where he recognized the hospital from the outside and would start crying hysterically.
After months and months we were still no closer to an answer. We've gone through therapies and medications like you wouldn't believe. We've had scares with his pancreas and liver and still nothing. I ended up nursing him until he was three just so he would have enough in him to survive. There was talks of a g tube if we couldn't get him eating.
Fortunately he did start to eat solids, to this day though he really struggles to eat. He can go more than half the day without eating and even then there are very few things we can actually get him to eat. His tummy issues are still there too. They seemed to get better for a while but lately it's been pretty bad. Tonight for example he's asleep next to me, tummy hard and bloated. Needed to go but can't or won't, maybe a bit of both. Some days like today he throws up because his tummy has no more room or maybe from the pain regardless it is awful. I wouldn't wish this stress on anyone.
It's not something I talk about all that often but it is definitely something I think about everyday. I worry he's not getting enough, I worry there is an underlying cause that we haven't yet discovered. My mind gets carried away to unimaginable fears.
Someday I hope we will get past this, I long for the days where I won't have to worry about dinner sending him into a panic because it's not one of the few foods he seems to tolerate. Until then we will just have to keep on pushing through no matter how exhausting it can be.


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With so many on offer these days, it is great to have a make you can trust. The Ride n Roll - Ladybird is certainly that and will be a excellent purchase. For this reduced price, the Ride n Roll - Ladybird comes highly respected and is always a regular choice for lots of people.
- Suitable from 12 months
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Product Description
A fun way for kids to get around. The base is made from top quality plywood with a tough rounded edge. The bodies are soft for a comfortable safe ride and the covers are made from vinyl that can be wiped clean with a damp cloth. Features soft grips and are omni-directional wheels, making the creature spin and twist for ultimate fun.


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@PowerScripting chat with vCO Managers
Did you miss the @PowerScripting folks as they interviewed the vCO Management Team?
If so, don't worry - the session was recorded! See below for details:
- Thomas Corfmat - vCO Product Manager
- Hemant Gaidhani - vCO Product Marketing Manager
with bonus commentary from one of VMware's PowerShell gurus: Alan Renouf
Learn more about vCO and the new support/integrations with PowerShell.
- PowerScripting Blog Article: http://powerscripting.wordpress.com/2012/01/16/episode-172-vcenter-orchestrator/
- Video: http://static.bambuser.com/r/player.swf?username=PowerScripting
- PodCast: http://feeds.feedburner.com/PowerScripting


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Paul Cohen earned his MD at Columbia University in 1981. He completed his residency in anatomic pathology at Massachusetts General Hospital and the National Cancer Institute, where he also completed a pathology fellowship. Dr. Cohen is board certified in anatomic pathology. He is the Chair of Pathology at Bridgeport Hospital, and an Assistant Professor of Pathology at Yale School of Medicine. Dr. Cohen is the Director of the Yale Pathology residency program at Bridgeport Hospital.
Paul Jay Cohen, MD
Pathology, Anatomic Pathology
Accepting new patients? Yes
Referrals required? From patients or physicians
Patient type treated: Child; Adult
Board Certified in Anatomic Pathology


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The Details for this job are shown below, if you would like more details or have any questions please use the contact details provided.
Freehand 2010 Limited is a small medical device company based in Guildford, Surrey. Our flagship product is a novel robotic system which holds and manipulates a laparoscope and camera during minimally-invasive (keyhole) surgery. We have recently broadened our design and development programme to cover devices in two different clinical areas and need an able Project Design Engineer to deliver these.
Product design - mostly mechanical some electrical
•SOLIDWORKS 3D CAD expertise•Ideally, experience in medical device product development •Some project management skills•Accuracy and thoroughness •Excellent verbal and written communication skills•Confidence to challenge assumptions to establish the correct product requirements
Salary Range:
Up to £38000
Guildford UK
Medical devices
Placement Type:
If you are interested in applying for this job or would like to find out more, please send CV along with introductory email identifying this position to the contact details below:


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The PSJA Parent Spotlight program features parents whose commitment, passion and dedication has motivated them to continue their education by participating in the Parental Engagement Program (PEP).
The program provides an opportunity to recognize parents of PSJA ISD students whose determination to succeed and improve their lives serve as inspiration and encouragement for their children and other parents in the PSJA community. Parents selected will be featured in the following outlets: district's website, social media platforms and KTRI Channel 17.
Parent participants must be nominated by parent educators/instructors or program administrators on the basis of their participation, achievements and development in the program.
To be considered, nominees must meet the following criteria:
- Be parents of at least one current PSJA student;
- Must be currently participating in the Parental Engagement Program (PEP);
- Must have been a participant of the Parental Engagement Program for at least one year prior to nomination;
- Must demonstrate outstanding leadership and commitment to continue their education.


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Man who abused young nieces has jail time reduced after appeal
The 48-year-old was convicted in 2018 of 25 counts of sexual assault.
Rapist (75) who abused sister and his seven daughters over 23-year period jailed for 20 years
From 1977 to 2000, James O'Reilly (75) subjected the young girls to "horrific" sexual abuse.
Harvey Weinstein reported to have tested positive for coronavirus
Sources told TMZ that Weinstein was one of two inmates to test positive at a New York prison.
'Depraved': Child rapist jailed for seven years
The survivor of the attack was the man's then-girlfriend's younger sister, who is waiving her anonymity.
One of Britain's most wanted fugitives convicted of murder following international manhunt
Shane O'Brien slashed 21-year-old Josh Hanson's neck at a west London bar in 2015.
Man with 56 convictions sentenced to four years in prison, two suspended, for dragging woman from car
He was found guilty of assault causing harm and engaging in threatening and abusive behaviour.
Six men escape death sentence for rape and murder of eight-year-old Indian girl
Investigations suggested that the girl was targeted in order to strike fear in her nomad community.
Man who was cleared of murdering former soldier is jailed for stabbing another man on same night
Gary Watson was convicted by a jury last February.
Man convicted of raping two women who were working as prostitutes
The man had pleaded not guilty to rape and sexual assault at a hotel in Portlaoise
Garda jailed for two years for possession of child abuse images
Joseph O'Connor (58) of west Dublin had pleaded not guilty.
Canadian teacher convicted after he used 'pen camera' to spy on female students
Ryan Jarvis (41) was charged with making video recordings of 27 female students.
Former senator found guilty of assaulting 3 gardaí
Heffernan appeared in court on charges brought following events relating to 31 July/ 1 August 2016.
Garda convicted of careless driving causing death of elderly woman
Warren Farrell was responding to a panic button call at a Topaz garage when the car struck Elizabeth Core.
Man convicted of €90,000 tiger kidnapping of postmistress, daughter and Italian student
Paschal Kelly (53) with an address at Cootehill, Co. Cavan, had pleaded not guilty to trespass and to false imprisonment.
Defence lawyer says Storm Emma looter was 'high on cocaine at the time'
David Berney received a two-year sentence today for burglary, but had 18 months suspended for the looting incident.
Convicted murderer who fled custody on Thursday arrested in Belfast
Samuel McKinley has fled from custody on more than one occasion.
Freddie Thompson found guilty of feud murder of David Douglas
He will receive the mandatory life sentence.
'I will never ever forgive you': Man to be sentenced in November for rape and sexual abuse of sisters
Bartholomew Prendergast pleaded guilty to two counts of rape and 13 counts of indecent assault of his sister-in-law Stephanie Hickey.
Man who raped sister-in-law when she was 15 is jailed for nine years
The woman described how she had no prior knowledge of sex the day the man raped her.
Man convicted of causing serious harm to two former partners by infecting them with HIV
The judge described the case as 'pretty difficult with unusual evidence'.
London man who made bombs for the Taliban convicted of terror offences
Ali was arrested by armed police near the Houses of Parliament in London on 27 April 2017.
Convicted of a €7.2 billion fraud - David Drumm was Anglo's man 'who called the shots' at the infamous bank
He will be sentenced on 20 June.
'Justice has been done': Victims and prosecutors react to Bill Cosby sex assault conviction
Cosby was convicted of drugging and molesting a university employee 14 years ago.
India approves death penalty for people convicted of raping children under 12
Violent crime against women has been on the rise in India despite tough laws enacted in 2013.
Jury convicts former New York nanny of murdering two children aged two and six
The 55-year-old is now likely to spend the rest of her life behind bars.
Man who exploited young girls ordered not to take part in computer programming course
Matthew Horan was sentenced to nine-and-a-half years with the final two years suspended.
British man given suspended sentence for Cambodia 'pornographic' pool party photos
Daniel Jones, 31, was among 10 foreigners arrested in January.
English club denies cover-up of Barry Bennell's sexual abuse and says it won't hold internal investigation
The former Crewe Alexandra youth coach was sentenced to 31 years in jail for 43 offences against young boys last month.
Former UK youth football coach Barry Bennell found guilty of child abuse
Bennell was convicted at Liverpool crown court on 36 counts of abuse between 1979 and 1990.
Irene White investigation turns to the person who paid Anthony Lambe to kill her
One avenue of investigation now being pursued by gardaí is that at least two other people were involved.
Two men, including one who has 25 wives and 146 children, convicted of polygamy
The men face up to five years in prison.
Convicted paedophile ordered to sell his house to pay €100,000 awarded to his nephew
The price will be negotiated in secret between John O'Neill and any prospective purchaser.
Fiancé of author Helen Bailey found guilty of murdering her and hiding body for 3 months
The court heard that he drugged Helen before killing her and hiding her body, he then sent her text messages and emails in a bid to cover his tracks.
Journalist convicted for bringing Syrian boy to Sweden
Fredrik Onnevall was filming a documentary when he met the 15-year-old boy in Greece.
Man who hoped his dog would get better without treatment convicted of animal cruelty
The man was in denial and hoped his pet would recover.
Australian firm responsible for Irishwoman's scalping injuries fined
Annie Dunne suffered the injuries while working on a farm.
Suspended jail sentences for four Chelsea fans over racist incident in Paris
Supporters prevented Souleymane Sylla from boarding the train on the night of a Champions League game in February 2015.
Man found guilty of murdering his two-month old daughter
Baby Daisy Mae Burrill suffered a large fracture to her skull and injuries consistent with having been shaken.
Serial killer Mark Nash loses appeal for damages
His lawyers claimed his human rights were breached by a delay in prosecuting him for the Grangegorman murders in 1997.
Irishman arrested in Spain after a year on the run
He had admitted supplying amphetamines but failed to appear in court for his sentencing in February last year.


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And of all the parlors of the world she walked into mine
Journeys. Life. Books and more.
Because real life gets boring And those characters keep bursting out. Of course Tyler has a life I need to write about 🙂
Funny Bone
Q: Why can't you tell a joke while standing on ice? A: Because it might crack up! You will not read jokes like these here. you will get worse 😀
It happens when you least expect it. Most regret it. And often when you never see it. Here's my own bit…
Fresh from the blog
An Uncommon Love that Saved me
One of the rarest gifts both my parents helped me achieve was extreme forms of self-confidence, which never bordered on foolhardiness. And yet, when I lost one of them to a sudden act of God, it was this self-confidence that took first hit. I had lost my mother to...
Why Freelancers & Self-employed need to Seek Community
As a self-employed individual, my life doesn't have HR managers or Finance teams to assist in my everyday activities. I rely on hiring designated firms or professionals for all the support functions. But more importantly I rely on a community of bloggers and social...
A Failure that Gave me Blogchatter
Very few people are aware about the life changing incident that I am about to share. It was the year 2012, I was newly married and floating on cloud nine of love and companionship. With that milestone achieved, I thought now could be a good time to push myself into...


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Amazing Escape Room
BC Events
Bliss Travels
Bowlero North Brunswick
Bury The Hatchet
Crossing Vineyards and Winery is tucked away on a 200-year old estate, conveniently located just off exit 10 of I-295 in the heart of Bucks County, Pennsylvania. This historic property offers a unique setting for weddings, rehearsal dinners, concerts, private parties and corporate gatherings.
Crossing is open from noon-6pm year-round and on Thursday evenings until 9pm. Join us for "Thursday After Hours" and enjoy a variety of fun classes and events. Hourly seated wine tastings at the Winery are offered on weekends and on holidays. Book on-line and save at www.crossingvineyards.com. Sample cocktails, beer and wine flights and Crossing Vineyards' award-winning wines by the glass or bottle. Light bites and table seating are also available. Enjoy vineyard views on the terrace and live music on selected Saturdays throughout the year.
Dave & Buster's
Despaña Princeton is an all-authentic Spanish (Spain) restaurant and tapas cafe with full service lunch and dinner. Despaña's unique space features a beautiful industrial chic second floor restaurant atmosphere with an outdoor terrace overlooking scenic Nassau Street. Guests can indulge after dining in the downstairs market and tapas café. The tapas café and market offers an exceptional menu for those on the go or for a quick café con leche or dessert. Despaña also offers full service catering and special event space and planning. Reservations recommended.
Consumers and designers who visit the Princeton Gallery will experience Miele's innovative, live product displays and may further their know-how through cooking programs and educational seminars. Miele's Gallery offers a fully functional kitchen area for classes, an elegant dining area for events and a coffee bar for customers to enjoy a perfect espresso or coffee while they visit. Classes and events for the design community and general public will be listed online at mieleusa.com/princeton.
The Gallery is part of Miele's total brand experience – communicating the quality, design and technology their products are known for around the world. From Prague to Hong Kong – Gütersloh, Germany (the location of Miele's global headquarters) to New York… the Princeton Gallery is among a growing list of successful Miele Gallery properties.
Pinot's Palette Princeton, an upscale paint and sip entertainment destination, was recently voted "Best Art Classes" by Town Topics Newspaper! Come find out why they are the best for a girls night out, date night, bachelorette party, corporate team-building, office party, or any special occasion. Anyone can be an artist and have fun being creative - no art experience required! In two or three hours you can create a festive, whimsical and colorful painting while sipping on your favorite beverage. With step-by-step instructions from one of the local artists, you will create the evening's painting in a fun and relaxed atmosphere! Pinot's Palette is BYOB and has partnerships with restaurants to offer a variety of catering options to make the most of your party! Pinot's Palette is conveniently located at the Princeton Forrestal Village right off US Highway 1.
Present Company
Great gatherings at Princeton Meadow Event Center! Every day, people come together to share and learn, to network, to perform and to celebrate. You supply the event-we provide an elegant setting with exceptional service and food.
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We are Sky Zone, creator of the world's first indoor trampoline park. Join us for a SAFE & FUN experience! We are an indoor trampoline park that not only features Trampolines but Warped Wall, Dodgeball, Toddler Zone, Free Climb Wall, Ninja Warrior Course, Basketball, and Foam Pit!
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The Grape Escape


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Tech Launch Arizona Can Help You Use Your Lab Materials to Drive Income
Campus Notes
Tech Launch Arizona Can Help You Use Your Lab Materials to Drive Income
You have unique, useful materials in your lab. Are they working as hard as they can for you?
Has your research demanded that you develop a new kind of material? Do you have something in your lab freezer that might be useful or valuable to other investigators? Do you receive requests for materials that your lab produces, such as:
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Research materials aren't typically patentable, but they certainly are valuable and can generate revenue.
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Our discussions are completely confidential, and we help you fulfill all your grant reporting requirements. Most research contracts contain language regarding reporting inventions; we'll help you understand your responsibilities.
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Thank you,
The Tech launch Arizona Team
PS. Want to share this information with your colleagues? We've made it easy for you; print the attached poster on 11x17 paper and post it in your lab!


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Uber, Careem drivers arrested as services remain suspended in Abu Dhabi
A source in Abu Dhabi said drivers had been detained over violations of regulations
As many as 50 drivers for Uber and Careem were arrested in Abu Dhabi before the ride-hailing companies withdrew services in the UAE capital on Saturday evening, according to local media reports.
The suspension of service remains in place, with one Abu Dhabi source familiar with the situation telling Reuters that some drivers had been detained over violations of regulations, but did not specify how many drivers or describe the violations.
Christian Eid, vice-president of marketing and communications for Careem, a Dubai-based company, said many of its drivers were being stopped by authorities in Abu Dhabi, apparently over licensing issues, and as a result had become nervous and were staying off the roads. This had forced Careem to halt services there, he said.
"A few of our cars have been stopped by authorities in the capital," said Eid, in a statement to Arabian Business.
"Until we have further clarification on the situation, Careem has decided to temporarily limit services in Abu Dhabi to avoid any inconvenience this may cause both our customers and captains."
The companies were attempting to resume operations as soon as possible, according to a spokesperson from Uber.
"As you may know, Uber and Careem decided to suspend their operations due to unforeseen circumstances," said the spokesperson.
"We want to let you know that this was a decision taken by both companies, and our goal is to resume operations in Abu Dhabi as soon as possible. Please note that Uber's services in Dubai are running as usual, we will have more information on Abu Dhabi in the next two days."
Arabian Business attempted to contact Abu Dhabi government's Centre for Regulation of Transport by Hire Cars, the Abu Dhabi taxi regulator, by both email and through social media, on Sunday but received no reply.
The centre oversees about seven taxi operators and 18 limousine operators, some of which are partly government-owned.
Despite rumours that services would also be suspended in Dubai, both firms insisted they would continue to pick up passengers in the emirate.
Uber, which launched services in Abu Dhabi in 2013, said last year that the Middle East and North Africa contained some of its fastest-growing markets and that it planned to invest $250 million to expand in the region.


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Ever since finding out about Frank Lee's Procession of the Ghouls back during Year 5 of this 32 Days of Halloween madness, I have been freaking obsessed. How I haven't managed to get up there and see it yet…no idea. We previously posted the mini-docu about the behind the scenes and origins and whatnot of this magnificent spectacle: a troupe of goblins and spectres of all sorts making their way through the Catheral Church of St. John the Divine in New York. It's sort of like a Krampus parade but indoors and if it had been conceived by the Henson Creature Workshop. Plus, afterwards they show a silent film. This year it's Hunchback of Notre Dame. (Seeing that in the fourth largest Christian church in the world…? On Halloween night! How cool would that be, right?)
This YouTube video isn't the best…but it will give you a taste of last year's Procession. For info on this year's, check out the official website here. And we posted a similar video for the 2012 Procession here.
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All about Coca-Cola Logos for your smartphone, your iPhone or any phone.A large collection of moving images of The Coca-Cola Company logo, one of the world's most popular logos.
Animated gifs and animations of Coca-Cola Logos to share on social networks, download or send to other phones for Whatsapp, Line or Skype. The Coca-Cola logo was designed by Frank Robinson, who also named the drink and drew the current logo with his handwriting. Although the drink was invented by pharmacist John Pemberton, were marketing techniques of Asa Griggs Candler entrepreneur who popularized the soft drink. The main version of the logo is white letters with red background. There are other versions, either for Coca-Cola Light, Coca-Cola Caffeine Free and Coca-Cola Zero, in which red, black, gray or gold colors, either for lyrics or used for background. The logo of the company has not changed much in its history.


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Surgeon and inventor Catherine Mohr tours the history of surgery (and its pre-painkiller, pre-antiseptic past), then demos some of the newest tools for surgery through tiny incisions, performed using nimble robot hands. Fascinating—but not for the squeamish.
IEET > Vision > HealthLongevity > Futurism > InnovationSurgery's Past, Present, and Robotic Future
Catherine Mohr Jul 20, 2011
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Urgent Care Doctor in Coral Springs, FL
If you've been looking for a top urgent care doctor in Coral Springs FL, then come see Dr.G at our Coral Springs urgent care clinic. We Are Open Late 7 Days a Week. We are Conveniently Located at 2224 N. University Drive Coral Springs, FL 33071. Call (561) 330-9363 For More Information!
For minor emergencies or urgent healthcare, come see an urgent care doctor! These professionals are available with no appointment necessary, unlike your family doctor who might not be available right away. Even the ER (emergency room) can be a hassle in some instances because you'll be put on a waiting list behind patients with major emergencies. Instead, come see our urgent care doctor in Coral Springs, FL. Our physician, Dr.Gs, would be happy to see you for any minor emergency or any other kind of common healthcare.
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In many instances, there are many advantages of choosing a walk-in and urgent care clinic over other options. One of the big advantages is that these clinics are open to walk-ins, so you don't have to make an appointment for care. You can simply walk right in for same-day service, which isn't an option with your family doctor. Walk-in and urgent care clinics are open extended hours too, just in case you need them after work or school.
Another big advantage with these clinics is that you can receive urgent treatment right on the spot. This means that for any minor emergency, like a minor sprain or strep throat, you can receive treatment within minutes of entering the clinic, with no wait times. For minor emergencies, you likely won't get immediate treatment at the ER.
There are many other advantages too, such as being able to take anyone in your family to these clinics, including children and seniors. Additionally, walk-in/urgent care clinics offer affordable services that are covered by insurance!
Is urgent care more expensive than a regular doctor?
Typically, urgent care is actually less expensive than going to see a regular doctor like your primary care physician. Regular doctors can cost hundreds of dollars before insurance coverage. Urgent care can cost half of the price before insurance coverage. Also, most urgent care providers work with most major insurance companies, so you can expect your services will be covered! Costs vary though, so to learn more about our costs here at Dr.Gs urgent care, get in touch with us at (561) 330-9363.
Some of the major providers we work with include, but aren't limited to, Blue Cross Blue Shield, Aetna, Humana, Tricare, United, Coventry, Medicare, Molina, Staywell, Cigna, workers comp, Avmed, Wellcare, and more. If you don't see your provider listed, that's okay, just call us to check whether we work with your insurance provider.
Who is Dr.G?
Dr.G is our owner and resident physician here at our clinic! Her full name is Dr. Mylissa Graber, MD, MBA, FACEP. The doctor is a board-certified physician with over 15 years of experience with minor emergency care. She also has a great deal of leadership experience, including being the former Executive Vice President of EmCare, and more. Today, she is dedicating herself to growing Dr.Gs Urgent Care and supporting you and your family. You can read more about Dr. Graber over on our Meet Our Team page.
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You can come right into Dr.Gs Urgent Care to meet with Dr.G or one of the professional physicians she trained herself! We have a Coral Springs location at 2224 N. University Drive Coral Springs, FL 33071. You can simply walk in for service without making any kind of appointment, and we're open for you Monday to Sunday, from 9 am to 9 pm.
We look forward to seeing you soon here at Dr.Gs Urgent Care!


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Would you recommend this product?
Whether it's for your LinkedIn, Facebook or Tinder ;-), an awesome Headshot with a professional photographer really ups your game. We have an fantastic studio where we offer hair and make up plus personal styling to help you look your best. Our goal is get you in and out as quick as possible while still giving you professional results. Standard Headshots at other places usually start at $200 and we start at $75 because we do all the shoots in one day. This helps keeps our costs low so we can pass the savings to you. In as little as 10 minutes you can have an awesome new look. We do ask you come a little early because SF traffic is notorious. But we try to keep the vibe pretty fun, we have some great music during your session and we offer snacks and mimosas to keep the energy high. Happy to answer any questions you may have!
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I love how you've complemented your Hireclub application/Facebook group (which focuses on getting face to face contacts) with a service that literally focuses on headshots. You're a clever man, and I appreciate you.
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Are you thinking of expanding to other cities ? I know a few photographers in LA that would eat this up! (Actors pay crazy prices for headshots) happy to try to connect them
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@swetzequity David our photographer is actually based in LA. We'd love to expand into as many cities as possible. Message me at email@example.com. Thank you!


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"Behind every great man there is a great woman."
I often heard people use this phrase to express about someone who is or had been very successful and powerful leader in the world. However, my story is different. Behind my simple story there are many great women: my late grandmothers, my mother, my aunts, the UNHCR and Red Cross female officers, my wife and recently my daughter who gave me the suggestion and idea for my book jacket. And this week many more of them have come to support me and listen to my story telling – behind me there are thirteen women and a giant-heart friend who is also there to support every charity event. I have come to know these people through my volunteer in one of the hardest hit towns, Ishinomaki, Miyagi Prefecture – approximately 400Km north of Tokyo, from the Japan 2011, 311 earthquakes.
I have also learned from my audience of the Nadia Book Club that many Japanese are familiar with the Second World War (WWII), and Vietnam War, but not too many people, especially the younger generation, know about the Killing Fields of Cambodia. Though it is not their fault, perhaps our history book categorises Cambodia Civil War from 1970 to 1975 as part of the Vietnam War. However, the audience sat there quietly clinging on to every single word of my untold story, Father Missed His Plane: A real-life story of a boy's separation from his family, survival and adversity in the Killing Fields of Cambodia and beyond. They were very keen to learn more from this very tragic event. Unlike the Australian audience I had back in early 1980s, once again it was not their fault either, it was simply at the time no one ever heard of the Khmer Rouge or believed that a little refugee boy like myself came from Cambodia and had gone through such a hardship in a concentration labour camp and survived. Yes, somehow, I survived, although I nearly didn't.
The recent UNHCR report shows more than half of the 5 million Syrian refugees are women and children under 17 years old. We must find a home for them to survive and for the children to get educated before they become an easy target for manipulation and turn themselves into those young soldiers of the Pol Pot's Khmer Rouge regime.
As you can see from the picture, behind 1 simple me, there are 13 great women (plus 1 great man).
This is an excerpt from Vincent Lee's untold story, Father Missed His Plane: A real-life story of a boy's separation from his family, survival and adversity in the Killing Fields of Cambodia and beyond. Read More


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More Details | Tray Identification Tag (110 x 38 mm) Single tag Latex free 15 colours
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To feel good and to be cheerful woman needs care and good mood. What can bring good mood? Its foot massage! and if this massage is made by your close loving person it has double effect. So let's practice it. The details of foot massage you can read in the next article >>
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