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Evolutionary change in response to natural selection will occur only if a trait confers a selective advantage and there is heritable variation. Positive connections between pollen traits and fitness have been found, but few studies of heritability have been conducted, and they have yielded conflicting results. To understand better the evolutionary significance of pollen competition and its potential role in sexual selection, the heritability of pollen tube-growth rate and the relationship between this trait and sporophytic offspring fitness were investigated in Collinsia heterophylla.
Because the question being asked was if female function benefited from obtaining genetically superior fathers by enhancing pollen competition, one-donor (per flower) crosses were used in order to exclude confounding effects of post-fertilization competition/allocation caused by multiple paternity. Each recipient plant was crossed with an average of five pollen donors. Pollen-tube growth rate and sporophytic traits were measured in both generations.
Pollen-tube growth rate in vitro differed among donors, and the differences were correlated with in vivo growth rate averaged over two to four maternal plants. Pollen-tube growth rate showed significant narrow-sense heritability and evolvability in a father–offspring regression. However, this pollen trait did not correlate significantly with sporophytic-offspring fitness.
These results suggest that pollen-tube growth rate can respond to selection via male function. The data presented here do not provide any support for the hypothesis that intense pollen competition enhances maternal plant fitness through increased paternity by higher-quality sporophytic fathers, although this advantage cannot be ruled out. These data are, however, consistent with the hypothesis that pollen competition is itself selectively advantageous, through both male and female function, by reducing the genetic load among successful gametophytic fathers (pollen), and reducing inbreeding depression associated with self–pollination in plants with mix-mating systems.
Over 80 % of flowering plants are hermaphrodites, meaning that both male and female reproductive functions may contribute to individual fitness (Richards, 1997). Despite this fact, selection pressures on the two reproductive functions can differ (Willson, 1979; Bell, 1985; Campbell, 1989; Maad and Alexandersson, 2004; Maad and Nilsson, 2004), potentially leading to sexual selection of floral traits (Skogsmyr and Lankinen, 2002; Delph and Ashman, 2006). Divergent selection pressures are likely to be particularly important during pollen competition in the pistil, because pollen has the opportunity to represent the sole 'interests' of male reproductive function. Although pollen performance directly reflects the male reproductive function, selection should favour female (pistil) reproductive traits that promote pollen competition, or otherwise screen pollen, since this may increase the fitness of her own offspring (Skogsmyr and Lankinen, 2002; Bernasconi et al., 2004).
Several studies have indeed shown that high pollen competitive ability, particularly pollen-tube growth rate, is associated with high siring ability (e.g. Snow and Spira, 1991, 1996; Marshall, 1998; Skogsmyr and Lankinen, 1999; Aronen et al., 2002). This indicates that selection on this trait may have evolutionary effects on the male function, provided that there is heritable variation for this trait (Fisher, 1958; Charlesworth and Charlesworth, 1992; Walsh and Charlesworth, 1992). Despite the great importance of heritability, there is only limited information available on levels of heritability of traits related to pollen competitive ability, and this only for a handful of species. Experiments with cultivated species suggest the presence of a genetic component of pollen competitive ability (e.g. Ottaviano et al., 1988; Schlichting et al., 1990; Sarkissian and Harder, 2001), but the few heritability studies in wild species have yielded contradictory results (see Snow and Mazer, 1988; Havens, 1994; Skogsmyr and Lankinen, 2000; Lamborn et al., 2005).
Since Mulcahy (1979) first suggested that pollen competition may be an important way to enhance offspring fitness by the screening of haploid pollen to eliminate malfunctioning genomes, numerous studies have investigated if enhanced pollen competition can improve offspring fitness. Assuming overlapping gene expression in gametophytes and sporophytes (Mulcahy et al., 1992, 1996; Walsh and Charlesworth, 1992; Hormaza and Herrero, 1994), older studies often found increased offspring fitness following higher levels of pollen competition (e.g. Mulcahy and Mulcahy, 1975; Winsor et al., 1987; Bertin, 1990; Quesada et al., 1993). These studies have, however, been criticized for not separating out the influences of male versus female reproductive functions (Charlesworth et al., 1987; Snow, 1994). More recent research has attempted to test mechanisms of how enhanced pollen competition and/or larger pollen loads can lead to higher offspring quality. They include (a) ovules being fertilized by highly competitive pollen produced by donors of superior genetic quality, (cf. 'good genes' of sexual selection theory; Zahavi, 1975; e.g. Skogsmyr and Lankinen, 2000; Pasonen et al., 2001), (b) larger pollen loads leading to increased sampling of superior, more-compatible, or genetically more variable pollen (e.g. Paschke et al., 2002; Bernasconi et al., 2003; Kron and Husband, 2006), and (c) maternal plants allocating more resources to ovules fertilized first or to ovules in pistils pollinated by more pollen (e.g. Delph et al., 1998). Other mechanisms include reduction of inbreeding depression by (d) self pollen performing worse than outcross pollen (cryptic self-incompatibility; e.g. Bertin and Sullivan, 1988; Kruszewski and Galloway, 2006) and (e) self pollen of low quality being avoided (Armbruster and Rogers, 2004; Lankinen and Armbruster, 2007).
The mechanism involving sexual selection (a above) in particular needs more research. For example, the relationship between offspring fitness and pollen traits of donors has been tested only in a few studies (see Mulcahy, 1971; Skogsmyr and Lankinen, 2000; Pasonen et al., 2001). These studies mainly used two-donor crosses, which indeed gives a direct test of pollen competitive ability. On the other hand, this approach may reduce the possibility of detecting paternal genetic effects on offspring quality because other factors can not be eliminated, such as the 'sampling effect' (Bernasconi et al., 2003) or differential allocation of maternal resources (Walsh and Charlesworth, 1992). When evaluating the relationship between offspring fitness and pollen competitive ability, it is also important to assess offspring fitness beyond the earliest offspring traits (cf. discussion on animals; Kokko et al., 2003). This is because traits expressed very early in the life-cycle of plants are known often to be strongly influenced by maternal effects (Roach and Wulff, 1987).
Previously it had been found that intense pollen competition in the self-compatible annual Collinsia heterophylla can reduce inbreeding depression following one-donor crosses of unequal pollen load size (Lankinen and Armbruster, 2007), suggesting a benefit of avoiding fertilization by self pollen of low quality. In order to understand better the possible selective advantages accruing from pollen competition, examined here are (a) the heritability and evolvability of pollen competitive ability, and (b) the relationship between pollen competitive ability and sporophytic fitness of donors and offspring at different stages of the life-cycle. To achieve this goal, one-donor crosses were performed on several recipients per donor in the greenhouse, in order to control for sampling effects and post-fertilization competition, as noted above. Because the aim was to study sexual selection of pollen competitive ability, rather than, for example, effects of genetic self-incompatibility, the focus was on evaluating the average performance of a donor across several recipients rather than estimating donor-by-recipient interactions (cf. Marshall, 1998). Even when donor-by-recipient interactions influence the success of pollen donors, selection will still favour a donor that is on average more successful than others. To assess pollen competitive ability, differences in pollen-tube growth rate were examined among the donors used in the crosses both in vitro (without any female influence) and in vivo. Also the performance of outcross and self pollen (in single pollinations) was compared in order to get an indirect test of whether more intense pollen competition would be advantageous as a means to avoid self-fertilization (cf. cryptic incompatibility, e.g. Bertin and Sullivan, 1988). It should be noted, however, that the test here is incomplete, as such differences are often only expressed when self and outcross pollen are growing together in the pistil (Aizen et al., 1990; Nemeth and Smith-Huerta, 2002; Kruszewski and Galloway, 2006).
Collinsia heterophylla (Plantaginaceae) is a self-compatible, diploid annual endemic to the California Floristic Province (Newsom, 1929; Neese, 1993). It is widely distributed and grows in shady places and on dry slopes (<1000 m). Flowering occurs between March and June depending on latitude and elevation. A variety of native bees serve as pollinators (Armbruster et al., 2002; W. S. Armbruster, unpubl. res.). Estimates of mean population outcrossing rates range from 0·32 to 0·64, based on allozyme markers (Charlesworth and Mayer, 1995), and up to 0·94 ± 0·27 (s.d.) based on morphological markers (Weil and Allard, 1964). Flowers are arranged in whorls on spikes. The zygomorphic flowers have five-lobed corollas comprising one upper and one lower lip. Corolla colour is generally white to pale purple on the upper lip and dark purple on the lower lip, although some populations can be pale purple or off-white on both lips. Flowers have four epipetalous stamens and one pistil. Newly opened flowers have undehisced anthers and non-receptive stigmas. As flowers develop, the anthers dehisce usually one at a time over the course of 3–4 d. During this period, the stigma becomes receptive and the style elongates. This eventually places the stigma in contact with the dehisced anthers, and self pollination can occur (for a more detailed description, see Armbruster et al., 2002; see also Kalisz et al., 1999). Four days after flower opening, the length of the style (including the stigma) is 13·2 ± 0·96 mm (± s.d.; n = 19; Å. Lankinen and W. S. Armbruster, unpubl. res.). Pollen is binucleate (Schrock and Palser, 1967). Ovaries develop into dry, dehiscent seed capsules.
A rough estimation indicates that at least 50 pollen grains can be in direct contact with the stigmatic surface at the same time (Å. Lankinen and W. S. Armbruster, unpubl. res), and the stigma can hold more pollen in additional layers. Because ovaries contain on average 11·2 ovules (max = 16 ovules; coastal populations; Armbruster et al., 2002; J. Maad and W. S. Armbruster, unpubl. res.), it is probable that the number of pollen grains on the stigma often greatly exceeds the number of ovules, suggesting that pollen competition can occur.
The approx. 40 maternal families used in this study had pale-purple flowers and originated from a population in Sisar Canyon, Ventura County, California (Population 4 in Armbruster et al., 2002). Plants were raised from seeds and grown in an insect-free greenhouse in winter/spring in a greenhouse at the Norwegian University of Science & Technology, Trondheim, during two seasons (parental and offspring generation). Because seeds from the different maternal families were not kept separate, the possibility cannot be excluded that some of the plants may have been full or half siblings, but this likely mimics roughly the structure of a field population in this self-compatible, mixed-mating species.
In order to evaluate heritability of pollen-tube growth rate and the relationship between pollen-tube growth rate and offspring fitness, a series of one-donor crosses was conducted. In total, 12 donors and 11 recipient plants were used. Some individuals were utilized as both donors and recipients. Pollen donors were combined in six pairs depending on their pollen-tube growth rate in a germination medium (see next section). Each pair consisted of one fast-growing and one slow-growing donor (the fast donors differed significantly from the slow ones, P < 0·0001). The reason for using this design was that we wanted to maximize the difference in pollen performance between the two donors crossed with each recipient plant. Seven recipient plants received pollen from two donor pairs, and four recipients received pollen from four donor pairs. Each donor was crossed with four to six recipients (mean = 5). All crossing combinations were repeated once. The total sample size was: 6 donor pairs × 2 donors per donor pair × 5 recipients/donor × 2 crosses per recipient = 120 crosses. Crosses involving the two donors in a pair were performed on different flowers of the same recipient plant only a few minutes apart on the same day. Crossing order of a donor pair in relation to other pairs was changed between recipients so that crossing order was as random as possible.
Crosses were performed on emasculated, fully receptive flowers (4 d after flower opening). Pollen from two flowers from the same donor plant was mixed on a microscopic slide. Pollen was added to the stigma directly from the slide until the stigma was completely covered with pollen. Crosses were repeated if the cross failed. This happened in only 3·8 % of all crosses. A few days before seed capsules were ripe, they were bagged to avoid losing any seeds.
Collected seeds were counted, weighed and eventually sown. Seeds from a cross were sown together (five seeds or less per pot) in ordinary potting compost. The pots were placed in a refrigerator (4 °C) for approx. 1 week to break seed dormancy. Pots were then placed in the greenhouse. Supplemental lights were on for 16 h a day. At the age of 4 weeks, six randomly chosen offspring per crossing combination (three per replicate cross) were repotted in one pot each and saved for assessment of later fitness components.
To investigate if pollen donors used in the crosses differed in pollen tube growth rate and to compare this trait in parents and their offspring, pollen was germinated in Hoekstra medium (Hoekstra and Bruinsma, 1975) in a dark chamber at a constant temperature of 20–21 °C. Pollen from two flowers per individual was sprinkled sparsely onto a drop of medium. In order to estimate in vitro pollen-tube growth rate, pollen tubes were allowed to grow in the medium for 1·5 h and the lengths of ten pollen tubes per sample measured under a light microscope (pollen tubes can be seen clearly without stain). Tubes of recently germinated pollen (shorter than 0·15 mm, i.e. less than approx. 25 % of average length) were not measured in order to reduce the influence of germination time on the metric. The average of the ten tubes was used in subsequent statistical analyses.
In vitro measurements can sometimes be unreliable, but an advantage is that pollen performance can be evaluated without influence of the maternal tissue. This is particularly important in a species with delayed stigma receptivity, where other options of assessing pollen performance excluding maternal influence are more difficult (e.g. bud pollination, Cruzan, 1993; see also Lyons et al., 1989). Because pollen tubes that were allowed to grow over night continued growing and became longer than the length of the pistil (Å. Lankinen and W. S. Armbruster, pers. obs.), it is assumed that pollen of C. heterophylla utilized both stored and exogenous resources (comparable to uptake of nutrients from the pistil after initial germination and growth; Herrero and Hormaza, 1996). To evaluate further the accuracy of the in vitro measurements using the parental generation, (a) repeatability was tested at the same greenhouse temperature, and (b) in vitro growth compared to pollen-tube growth rate in receptive pistils (see below). In the parental generation, pollen from all donors was germinated at the same time to control for temperature differences (see below; conducted twice, the average of the two measurements being used in correlation tests).
Simultaneous germination of pollen was not possible in the offspring generation, however, because of the large number of offspring analysed. Instead, pollen was germinated in 13 different batches, each consisting of 25 samples (representing 25 individual plants). Three of the 25 individuals per batch were germinated in two additional germination batches. Using the mean of the repeatedly germinated donors made it possible to standardize values in order to make all 13 batches comparable to each other. This standardization worked well because donors did not vary markedly in their response to temperature (ANOVA interaction term not significant; performance at 25 °C versus approx. 20 °C was 0·505 versus 0·333 mm 1·5 h−1, respectively; two-way ANOVA (mixed model); temperature: F1,11·7 = 41·8, P < 0·001; pollen donor (random factor): F17,6·55 = 5·08, P = 0·021; interaction: F11,16 = 1,00, P = 0·485; Å. Lankinen and W. S. Armbruster, unpubl. res.). Pollen germination in the offspring generation was conducted over a period of 2 weeks. Individuals were assessed once in the order in which they started flowering (approx. 1–2 weeks after they started flowering).
To assess pollen performance in the pistil, pollen traits were evaluated in fully receptive pistils (i.e. 4 d after flower opening) across 20 pollen donors in the parental generation (including the 12 donors used in the crosses). Pistils were collected after 1·5 and 2 h, respectively, because it takes about 1–1·5 h or longer for pollen to germinate on the stigma (Å. Lankinen, pers. obs.). Because it is crucial to assess performance averaged over several pistils to be able to tell if in vitro measurements reflect performance in receptive pistils, pollen from each of the 20 donors was placed on stigmas of emasculated flowers on three (or more) of the 13 recipient plants. To compare pollen-tube growth rate of self and outcross pollen, self pollen was placed on one stigma of each pollen donor. Crosses were distributed fairly evenly across the recipients receiving outcross pollen (between two and nine donors per recipient). In a few cases the same crossing combination was replicated, and the average was used. Minimum sample size = 20 pollen donors × 4 recipients per donor = 80 crosses for each of the two collection times. Some data were missing for pollen-tube growth rate, as a result of excluding data points when there was only one of the two measurements of pollen tube length (after 1·5 and 2 h) and when both measurements were zero. Sample size for pollen tube growth rate was reduced to 20 pollen donors × 3·35 recipients per donor = 67 crosses.
Collected pistils were immediately stored in ethanol (70 %). To stain pollen tubes, pistils were rinsed with distilled water and tissues were softened in 1 m NaOH for 2–3 h. Pistils were then thoroughly rinsed under distilled water and stained in a dye solution of aniline blue in aqueous K3PO4 for at least 3 h.
The length of the longest pollen tube was measured under UV epifluorescence light. The maximum rather than the mean pollen-tube length per sample was used in this experiment, because it was only possible to distinguish tissue with or without pollen tubes (though it was fairly easy to determine the tip of the longest pollen tube). The in vivo value that was compared with the in vitro measurement was thus the mean value of the longest tube seen in each of several recipients. Even though in vitro and in vivo measurements are slightly different, a positive correlation between the two measurements would still indicate that tube growth rate in the artificial medium reflects how far down the style pollen-tubes will have grown. Pollen-tube growth rate was estimated as the difference in pollen tube length between 2 and 1·5 h.
Sporophytic traits were measured in donors involved in one-donor crosses and in the resulting offspring. For the parental generation, seedling height was measured at 9 weeks, day of first flowering, and final plant height. In the offspring generation, seeds were counted and weighed to get an estimate of mean seed weight of the seeds that became the offspring generation. Total germination percentage was recorded, and seedling size measured at both 4 and 7 weeks. Early seedling size was estimated by measuring the width of the entire seedling (most seedlings only consisted of two pairs of leaves). Later seedling size was measured as the plant height times the total number of branches. In the year of the offspring generation, plant growth rate was much faster (probably because plants were grown later in the spring, hence experienced longer days). Seedling size at 9 weeks in the parents thus equalled the size somewhere between week 4 and week 7 in the offspring. The date of first flowering was noted. Number of flowers in the first inflorescence (main shoot) was measured as an indication of plant size. Seed production was investigated by counting seeds in three capsules per individual. Even though all seeds produced by an individual were not counted, it is likely that the two traits assessed, number of flowers on the main shoot and seeds per capsule, strongly influence life-time seed production.
When testing the relationship between in vitro and in vivo pollen-tube growth rate, data were pooled from related and unrelated pistils in order to get as large a sample size as possible (because of the large measurement error in the pistil). No differences were found between pollen types; see Results. Even though the distribution of in vivo pollen-tube growth rate did not deviate from normal, the distribution of the error terms did. However, because the distribution of error terms was not skewed, this deviation should not be a serious problem (Sokal and Rohlf, 1995). It was not possible to improve the distribution by transformation, nor was an appropriate non-parametric test found.
Narrow-sense heritability (h2) of pollen-tube growth rate was estimated in vitro in a father–offspring regression (Falconer and McKay, 1996), using a hierarchical analysis [a nested ANOVA that takes differences in sample size (of paternal offspring groups) into account; Sokal and Rohlf, 1995]. The level of replication used to test the regression is not different from that in a more ordinary test involving the unweighted mean of each paternal offspring group. Because heritability may be an inaccurate assessment of the ability of a trait to evolve (for a detailed discussion see, for example, Houle, 1992; Hansen et al., 2003), evolvability (IA) was also estimated as the additive genetic variance (VA) scaled by the square of the trait mean (Z): IA = 100 × VA/Z2 = 100 × h2 × σ2T/Z2 (Hansen et al., 2003), where σ2T represents the total phenotypic variance.
When analysing relationships between pollen-tube growth rate of donors and quality of donors or offspring, we used pollen-tube growth rate in vivo rather than in vitro. The relationship between in vivo pollen-tube growth rate of donors and sporophytic quality of their offspring was evaluated using the same kind of hierarchical analysis as for the heritability estimates of pollen-tube growth rate (see above).
Differences were analysed between self- and outcross-pollen performance after 1·5 and 2 h with a mixed model, using pollination type (self or outcross) as a fixed factor and pollen donor and the interaction as random factors. Because the error terms deviated significantly from normality, the difference between self- and outcross-pollen performance was also tested with a non-parametric test (Wilcoxon signed rank test). This test showed the same result as the parametric test (P > 0·20 for both traits).
Most analyses were performed with SPSS (2004), specifically the GLM procedure for the ANOVAs. The GLM procedure uses weighted least squares to estimate model parameters and can deal with an unbalanced design. The Satterthwaite method was used to calculate the degrees of freedom in the mixed models. Type-III sums of squares were used in all ANOVAs.
Pollen started germinating within 15 min in the germination medium. Germination ability was generally high (>90 %). Pollen donors differed in pollen-tube growth rate in the medium (one-way ANOVA; F11,23 = 6·63, P = 0·0014). The donor with fastest growing pollen was on average 2·14 faster than the slowest donor. The repeatability of pollen-tube growth rate was high (strong correlation) in the same greenhouse temperature (Pearson correlation; r = 0·853, d.f. = 9, P = 0·008), suggesting high consistency of in vitro measurements.
Pollen from all donors had initiated pollen-tube growth in a majority of the pistils (79 %) after 1·5 h [mean tube-length 2·2 ± 0·86 (s.d.) mm]. At 2 h the number of pistils with pollen tubes present had not increased, but pollen tubes were significantly longer [mean tube-length 2·8 ± 1·6 (s.d.) mm; paired t-test; n = 20, P = 0·02]. Pollen-tube growth rate (difference in tube-length between 2 h and 1·5 h) showed large variation [mean 0·7 ± 2·1 (s.d.) mm 0·5 h−1], probably partly as a consequence of a large measurement error caused by the fact that different pistils were collected at 1·5 h and 2 h and hence were independent. The difference in tube-length between 2 h and 1·5 h ranged from negative to positive values, which means that the mean value may be unreliable (probably an underestimate). Growth rate of the fastest donor was 3·35 mm 0·5 h−1, but it was not possible to get a reliable estimate of the absolute performance of the slowest donor as the slowest donor could not have had a negative value. The fastest pollen donor should grow to the base of the style (13 mm) in 3–3·5 h (including 1–1·5 h of germination), assuming growth rate is constant through the style. Preliminary data indicated that 46 % of pistils had pollen tubes at the base of the style 3 h after pollination and 93 % had pollen tubes at the base after 4 h (Å. Lankinen, W. S. Armbruster, unpubl. res.). It is thus probable that tube growth rate is slightly higher nearer the base of the style.
Despite the large variation in the in vivo estimates, the pollen-tube growth rates in germination medium and in pistils were correlated (Pearson correlation; r = 0·448, d.f. = 18, P = 0·048), suggesting that growth rate in medium reflects the average growth rate in the 3·35 recipient plants per donor. This result also indicates probable in vivo differences among donors. There was an even stronger relationship between pollen-tube growth rates of the fastest tube in the medium and in vivo measurements (Pearson correlation; r = 0·575, d.f. = 18, P = 0·008), presumably reflecting that this in vitro measurement was more similar to what was actually measured in the pistil. In the medium, measurements of the longest tube and the average of ten tubes were strongly correlated (Pearson correlation; r = 0·931, d.f. = 18, P << 0·0001). Pollen-tube growth rate was on average 3·37 times faster in pistils than in medium (paired t-test; n = 20, P = 0·11).
The father–offspring regression coefficient for pollen-tube growth rate in vitro was significant (Fig. 1). This indicates that there is additive genetic variation for this trait (narrow-sense h2 (± s.e.) = 0·356 ± 0·145; P = 0·018; n = 12 fathers, 221 offspring).
Evolvability, IA, of pollen-tube growth rate was calculated to be 3·8 % (IA = 100 × VA /Z2 = 100 × h2 × σ2T/Z2 = 100 × 0·356 × 0·02372/0·4712). The regression between mothers and offspring did not, however, have a significant slope [hierarchical analysis; h2 = 2 × b = 0·161 ± 0·234 (s.e.), P = 0·50; nmaternal offspring groups = 11, noffspring = 221], suggesting maternal effects cancelled out or swamped additive genetic effects.
No significant relationships between pollen-tube growth rate in pistils and sporophytic traits were detected in pollen donors of the parental generation (Table 1). Similarly, no significant correlations were detected between pollen-tube growth rate in vitro and sporophytic traits of individuals in the offspring generation (Table 2).
Between generations, pollen-tube growth rate was significantly correlated with only one offspring trait: there was a negative correlation with the earliest measurement of seedling size (Table 3). This correlation was not significant, however, after correcting for multiple comparisons.
Pollen tubes were present more often in pistils pollinated with self pollen than pistils pollinated with outcross pollen (Wilcoxon signed ranks test: self, 25 out of 26; outcross, 99 out of 134; T+ = 46, n = 10, P = 0·032). Likewise, pollen was better at germinating on pistils of the donor individual ('self pistils') than on unrelated pistils (Wilcoxon signed ranks test: self, 42 out of 46; outcross, 108 out of 144; T+ = 115·5, n = 17, P = 0·010). Pollen-tube growth rate did not differ significantly between pollination types (self vs. cross): mean unrelated (± s.d.) = 0·78 (± 1·7); mean self (± s.d.) = 0·72 (± 2·0) mm (Table 4).
Pollen-tube growth rate in vitro differed among pollen donors in Collinsia heterophylla. This gametophytic (pollen) trait also correlated with average in vivo growth rates on the two to four flowers receiving the pollen. Narrow-sense heritability and evolvability of pollen-tube growth rate in vitro were significant, as calculated by father–offspring regression. However, no positive relationships between pollen-tube growth rate and sporophytic fitness was found in either the parental or offspring generation.
Pollen traits can vary among pollen donors, and such variation can generate differences in siring success (e.g. Snow and Spira, 1996; Skogsmyr and Lankinen, 1999). In some species, however, recipient plants influence pollen competitive ability so that the rank order of donors is inconsistent across recipients (sire–dam interaction); this is often connected to self-incompatibility, inbreeding-depression or outbreeding depression (e.g. Fenster and Sork, 1988; Cruzan, 1990; Johnston, 1993; but see Marshall, 1998). A pollen trait can, however, be selected for even when there is female influence, provided the trait confers a high average pollen competitive ability. In this study of C. heterophylla, measurement error of pollen-tube growth rate in the pistil was large. This was presumably at least partly caused by the large variation in pollen-germination time in the pistil in combination with the use of two different pistils (collected at two time intervals) to assess pollen tube growth rate. Nevertheless, in vitro pollen-tube growth rate of donors correlated with average in vivo growth rate across multiple recipients. This suggests that the variation in pollen tube growth rate among donors was on average consistent across maternal plants.
Finding large variation in pollen-germination time may indicate that the ability to germinate fast may be more important for siring success than inherent differences in pollen tube growth rate. A numerical comparison of the average length of fully receptive stigmas and styles with the maximum difference in pollen tube growth rate between donors in C. heterophylla (fast/slow = 2·14, provided the in vitro ratio was similar to the in vivo ratio), indicated that the fastest donor will reach the base of the style 2·2 h sooner than the slowest donor [2 h (fast)–4·2 h (slow)]. Thus the variation among donors in when pollen tubes reach ovules should be considerably greater than the variation in germination time (maximum is approx. 0·5 h). It should thus be possible for fast-growing donors to overtake slow-growing ones, at least when donors are forced to start germinating on the stigma at a similar time. Because stigma receptivity is delayed in C. heterophylla and an unreceptive stigma can collect pollen for several days (Armbruster et al., 2002; Lankinen et al., 2007), it is likely that arrival time of pollen has little effect on reproductive success.
Heritable variation of pollen performance was not detected in Raphanus raphanistrum (Snow and Mazer, 1988). Similarly in Oenothera organensis, only a small portion (9 %) of the variation in pollen tube growth rate was explained by the genetic component (Havens, 1994). More recent studies have, however, detected higher heritabilities of pollen traits likely associated with performance: pollen-tube growth rate (49 % in Viola tricolor; Skogsmyr and Lankinen, 2000), pollen-grain size (19–40 % across three populations of Mimulus guttatus; Lamborn et al., 2005), and (probably) pollen-germination rate (36 % in intra- and inter-population crosses across six populations of Silene latifolia; Jolivet and Bernasconi, 2007). It is, of course, not certain that the last two traits are related to pollen competitive ability. The present estimate in C. heterophylla indicates that narrow-sense heritability of pollen tube growth rate was approx. 36 %, a value within the above range. The present estimate of evolvability indicates that the trait will change 3·8 % per generation in response to a standardized selection gradient of 1 (i.e. selection as strong as selection on fitness itself; see Hansen et al., 2003). In comparison, evolvabilities ranged between 0·01 % and 1·71 % for 24 floral traits in Dalechampia (Hansen et al., 2003) and between 0·16 % and 2·24 % for 11 floral characters in Campanula rotundifolia (J. Maad and W. S. Armbruster, unpubl. res.). The present value was surprisingly high, given that pollen-tube growth rate has been expected to have little evolutionary potential (e.g. Charlesworth and Charlesworth, 1992), in comparison to floral morphological traits.
Because temperature affects pollen-tube growth rate in C. heterophylla, and because only one (standard) temperature was considered in the present study, the estimate of heritability might be higher (or lower) than under field conditions. It seems unlikely, however, that the evolvability is overestimated for this reason, because this estimate of evolvability is expected to be largely unaffected by environmental variance (Houle, 1992; Hansen et al., 2003). In future studies, it will be important to confirm that measurements of pollen-tube growth rate are related to siring ability in this species, as in some others (e.g. Snow and Spira, 1991, 1996; Marshall, 1998; Skogsmyr and Lankinen, 1999; Aronen et al., 2002), because selection only acts on a heritable trait if it confers higher survival or reproductive success.
Delayed stigma receptivity may enhance pollen competition in C. heterophylla (Armbruster et al., 2002; Lankinen et al., 2007; see Galen et al., 1986). This raises the question of what potential benefits may accrue from more intense pollen competition. A previous study showed that a possible advantage of pollen competition in this self-compatible species is reduced inbreeding depression (Lankinen and Armbruster, 2007; see also Armbruster and Rogers, 2004). The current study was designed to test the potential benefit of higher offspring fitness when fertilization is by pollen from sporophytes with more competitive pollen (see Mulcahy, 1979). If it is advantageous to be fertilized by pollen that is more competitive, relationships between pollen competitive ability and vigour, both within and between generations, would be expected (cf. good-genes hypothesis; Zahavi, 1975; see reviews by Andersson, 1994; Skogsmyr and Lankinen, 2002). In inbred lines of Zea mays, Mulcahy (1971) found a positive correlation between relative pollen-tube growth rate of two competitors and relative seed weight in the next generation. Because one-donor crosses did not result in the same correlation and because seed weight is not usually influenced by paternal genotype, competitive interactions between developing seeds probably caused the positive correlation. Such effects may or may not be connected with higher offspring fitness. Following one-donor crosses in Betula pendula, donors with higher pollen-tube growth rate sired offspring with heavier seeds, but not larger seedlings (Pasonen et al., 2001). In Viola tricolor, on the other hand, more competitive pollen gave rise to offspring that themselves produced more seeds (Skogsmyr and Lankinen, 2000). No support was found in C. heterophylla for the predictions that donors with faster-growing pollen have greater sporophytic vigour or that their sporophytic offspring show higher fitness in any part of the life-cycle. The power (sample size) could, however, have been too low to detect real relationships, or it is possible that differences in offspring quality were not expressed in greenhouse conditions (see Kalla and Ashman, 2002). Maternal effects (Roach and Wulff, 1987) or effects of maternal condition could also have masked relationships between generations. In Raphanus sativus, for example, pollen donors with relatively higher pollen-competitive ability on young maternal plants produced earlier-maturing offspring, but this relationship was not seen on older maternal plants (Marshall et al., 2007).
The only apparent benefit of being fathered by donors with high pollen-tube growth rates detected in C. heterophylla is that the offspring themselves also have high pollen-tube growth rates. It is worth noting, as has been pointed out recently, that a negative correlation between reproductive success and viability may result if 'male attractiveness' is increased at the expense of viability (Kokko, 2001). In this case a negative correlation between reproductive success and viability may still be connected to higher total offspring fitness (including offspring fitness through male reproductive success). Interestingly, in another study on C. heterophylla, pollen donor identity affected how early during floral development seeds could be sired (Lankinen and Kiboi, 2007). Pollen donors that were able to fertilize seeds early had higher germination speed, possibly indicating that competitive ability of donors differ depending on how receptive the stigma is. In addition to investigating how pollen traits are related to siring success on fully receptive pistils, in the future it would also be interesting to study what determines pollen competitive ability at earlier stages of floral development.
A further partial test of an additional potential benefit of enhanced pollen competition was made: cryptic self-incompatibility, leading to a higher proportion of out-crossed offspring (see Bertin and Sullivan, 1988). However, the present results suggested that cross-pollen does not perform better than self-pollen in the pistil. (Indeed self-pollen can sometimes perform better than outcross pollen, based on germination percentage on fully receptive stigmas.) On the other hand, the limitations of the present in vivo measurements (it was only possible to measure the longest pollen tube) made it hard to compare average pollen tube lengths of the two pollen types. It is also important to note that only the outcome for the two pollen types in separate pistils was investigated. It remains a distinct possibility that outcross pollen performs better only when both pollen types are growing together in the pistil (Aizen et al., 1990; Nemeth and Smith-Huerta, 2002; Kruszewski and Galloway, 2006). Because C. heterophylla has a delayed selfing mechanism (Armbruster et al., 2002), it is possible that selfing mostly occurs when pollinators failed to visit, i.e. selfing will mainly be beneficial in terms of reproductive assurance. This could potentially lead to weak selection for pistil mechanisms that reduce the performance of self pollen.
Provided that pollen-tube growth rate influences pollen competitive ability in Collinsia heterophylla as in other plants, the degree of phenotypic and heritable variation reported here indicates that this trait can respond to sexual selection on the male reproductive function. The present data do not, however, suggest an advantage for the female reproductive function of favouring sporophytic fathers with fast-growing tubes, apart from the gametophytic benefit in the next generation (i.e. offspring that themselves produce pollen with faster-growing tubes). The present data, though incomplete, suggest an absence of cryptic self-incompatibility in this species and that the benefit of promoting pollen competition is unrelated to the ability to favour outcross pollen over self pollen. However, studies comparing self and outcross pollen growing simultaneously in the same pistils are necessary before this possible benefit can be ruled out. It is important to note that the present results do not exclude the possibility that pollen competition generates an advantage to female function in terms of higher offspring quality. If the pollen load is small, ovules are likely to be fertilized by lower-quality pollen (independent of pollen donor quality). Thus features that increase the size of the pollen load should increase female reproductive fitness, as has been shown in numerous previous studies (e.g. Johannsson and Stephenson, 1997; Quesada et al., 2001; Davis, 2004). The fitness advantage of larger pollen loads may be even stronger when the pollen load only consists of self pollen (Armbruster and Rogers, 2004). Indeed, the only detected advantage of enhanced pollen competition in C. heterophylla thus far, is the ability to reduce inbreeding depression (by increasing the relative quality of selfed offspring), as revealed by varying the size and composition of the pollen load (Lankinen and Armbruster, 2007). In future studies, it will be important to assess whether the main advantage of large pollen loads is a larger sample of superior donors and/or genetically more diverse pollen (Bernasconi et al., 2003; Kron and Husband, 2006) rather than more intense pollen competition per se (but see Armbruster and Rogers, 2004).
We wish to thank A. Stephenson and an anonymous reviewer for useful comments on the text. This work was supported by European Science Foundation (Plant Adaptation programme); the Knut and Alice Wallenberg Foundation; Marie Curie Individual Fellowship; the Norwegian Research Council; the US National Science Foundation (grant number DEB-0324808), the Royal Swedish Academy of Sciences and the Swedish Research Council.
PubMed Central Canada is a service of the Canadian Institutes of Health Research (CIHR) working in partnership with the National Research Council's national science library in cooperation with the National Center for Biotechnology Information at the U.S. National Library of Medicine(NCBI/NLM). It includes content provided to the PubMed Central International archive by participating publishers.


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xpswmm & xpstorm 2017
Now Available in the U.S. and Canada
Introducing xpswmm and xpstorm 2017, the custom built integrated 1D/2D Modeling System by XP Solutions. Watch this premiere release video to see how xpswmm can streamline your workflow and make your modeling work more efficient.
Free Webinar: Build a 1D/2D Model from Scratch
This highly requested webinar by Anthony Kuch, MSc, will assist those who want to see how a new 1D/2D flood model is built. PDH credits available to those who register and attend!
Free Webinar: Preparing Data for Sanitary Sewer Modeling
Preparing long term maintenance plans for a sanitary or combined sewer system involves a lot of moving parts. Flows during dry weather and wet weather must be understood to insure that maintenance/repair/replace dollars are targeted. Modeling Rainfall Derived Inflow and Infiltration using the RTK method helps us understand total demand on our sanitary systems. This webinar will discuss the data requirements for using these techniques, and the importance of adequately preparing the inputs used for these methods.
Latest News
Every Drop Counts - A Call to Innovation
by Zach Sample, P.E.
Stormwater Products Manager
A Request for Startups post on January 3rd on the Y Combinator Blog caught my eye. The blogger talked about the need to prepare for things to get worse with regard to climate change, and called for applications for funding from those working...
Get More Out of Water Resources Engineering Models
by Venu Kandiah, PhD, P.E.
Senior Water Resources Engineer
The effort required to develop water resources engineering models can be significant. But once developed, in many instances the models are only evaluated for a limited, or a single set of conditions. The results and predictions obtained from these evaluations are...
How to Get From Here to There - Professional Training
by Anthony Kuch, MSc
VP Client Services
On a recent flight to instruct at one of our xpswmm/xpstorm training workshops that we hold around the country, I had the opportunity to connect on the airline's Wi-Fi. As we crossed the country I could see that we would hit...
Free Webinar: Building a 1D/2D Model from Scratch
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Wed 01 Mar 2017 at 08:00 PST
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(Wed 15 Mar 2017 at 11:00 EDT)
Training in St. Louis, MO: Stormwater, Sanitary and Flood Modeling
St. Louis, MO, United States
Wed 22 Mar - Fri 24 Mar 2017

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Get the highest video recording performance with Class 10 rating for Full HD video (1080p), plus more speed to catch the shot with your point and shoot camera. With SanDisk Ultra UHS-I cards you'll...More
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Specialty Food
- Vegetables and Tomatoes
- Speciality Vegetarian Dishes
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Organic American Veggie Burger Product ID: NFF974097 Amys
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Product Id: NFF974097
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DescriptionShips Frozen Extra shipping appliesAmysMfg#: 00015
Ingredients : (Vegan) Organic Onions, Organic Mushrooms, Filtered Water, Wheat Gluten, Textured Soy Protein Concentrate, Organic Bulger Wheat, Organic Celery, Organic Carrots, Organic Walnuts, Organic Oats, Organic Tomatoes, Organic Bell Peppers, Expeller Pressed High Oleic Safflower And/Or Sunflower Oil, Organic Sweet Rice Flour, Spices, Sea Salt, Organic Potatoes, Organic Garlic, Organic Evaporated Cane Juice, Vinegar, Organic Molasses, Natural Hickory Smoke Flavor, Ground Mustard Seeds. Contains Wheat, Soy And Walnuts
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Reducing Antibiotic Prescriptions »
"A unique type of poster placed in exam rooms helped reduce unnecessary antibiotic prescriptions for respiratory infections during flu season. The approach could help reduce costs and extend the usefulness of these drugs.
Antibiotics a"...
Patients should be informed that Mellaril (thioridazine hcl) has been associated with potentially fatal heart rhythm disturbances. The risk of such events may be increased when certain drugs are given together with Mellaril (thioridazine hcl) . Therefore, patients should inform the prescriber that they are receiving Mellaril (thioridazine hcl) treatment before taking any new medication.
Given the likelihood that some patients exposed chronically to neuroleptics will develop tardive dyskinesia, it is advised that all patients in whom chronic use is contemplated be given, if possible, full information about this risk. The decision to inform patients and/or their guardians must obviously take into account the clinical circumstances and the competency of the patient to understand the information provided.
Last reviewed on RxList: 3/31/2009
This monograph has been modified to include the generic and brand name in many instances.
Mellaril FDA Prescribing Information
Additional Mellaril Information
Mellaril - User Reviews
Mellaril User Reviews
Now you can gain knowledge and insight about a drug treatment with Patient Discussions.
Here is a collection of user reviews for the medication Mellaril sorted by most helpful.
Report Problems to the Food and Drug Administration
You are encouraged to report negative side effects of prescription drugs to the FDA. Visit the FDA MedWatch website or call 1-800-FDA-1088.
Women's Health
Find out what women really need.


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Phone: (289) 320-8250
1238 Dominion Road
Fort Erie, Ontario, L2A 1H7
Slinger Football Dog Toy - 7"Play fetch for hours with the Nerf Slinger Football. This dog toy has an integrated rope handle with knotted end so you can easily launch it far distances. It's made of durable nylon with triple stitching for superior durability.
Ecological Attributes
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This product is eco-responsible because it reduces the impacts associated with the following environmental issues:
Click on the icons
to see why this product is eco-responsible
There are no significant environmental benefits at this stage in the life cycle.
There are no significant environmental benefits at this stage in the life cycle.
1238 Dominion Road, Fort Erie,
Ontario, L2A 1H7
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created by PomPom Games and published by PomPom Games, Crystal Interactive, Inc. & Mastertronic Ltd.
Classic shoot 'em up action as pure as you can get. Space Tripper dares you to be retro and reminds us all of what gaming should be about!
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Kinase inhibition has emerged as the preeminent point of intervention for developing novel anti-cancer therapeutics and research tool compounds to dissect signaling pathways. In this proposal, we focus on redox- sensitive kinases and the development of inhibitors that target the oxidized form of these proteins, which are prevalent in many human cancers.
In Aim 1, we identify chemical warheads that target cysteine oxidation species.
In Aim 2, we develop molecules that target oxidized tyrosine kinases and evaluate their effectiveness in vitro and cancer cell-based assays.
In Aim 3, we evaluate the therapeutic efficacy of new kinase inhibitors. Inhibitors identified in this study will serve as vital tools for dissecting mechanistic details of kinase redox regulation and the relationship between oxidative stress/aberrant kinase activity in cancer. Furthermore, these collective aims will create a foundation for the development of an entirely new class of anti-cancer drugs.
Oxidative stress and deregulation of protein kinase activity play significant roles in the development and progression of cancer. Successful covalent inhibitors have been developed that interact with the amino acids cysteine or lysine in proteins, however off-target interactions remain a significant problem. Here, we propose a new chemical strategy to make inhibitors that retains the specific advantages gained through covalent targeting and decrease the potential for off-target activity. These new inhibitors will provide an important starting point for the development of new anti-cancer drugs.
|Yang, Jing; Gupta, Vinayak; Carroll, Kate S et al. (2014) Site-specific mapping and quantification of protein S-sulphenylation in cells. Nat Commun 5:4776
|Martínez-Acedo, Pablo; Gupta, Vinayak; Carroll, Kate S (2014) Proteomic analysis of peptides tagged with dimedone and related probes. J Mass Spectrom 49:257-65
|Zhang, Dehui; Macinkovic, Igor; Devarie-Baez, Nelmi O et al. (2014) Detection of protein S-sulfhydration by a tag-switch technique. Angew Chem Int Ed Engl 53:575-81
|Gupta, Vinayak; Carroll, Kate S (2014) Sulfenic acid chemistry, detection and cellular lifetime. Biochim Biophys Acta 1840:847-75
|Garcia, Francisco J; Carroll, Kate S (2014) Redox-based probes as tools to monitor oxidized protein tyrosine phosphatases in living cells. Eur J Med Chem 88:28-33


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Canon BP-514 Battery
Guide price: £48
Last updated on 30th Aug 2013
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Canon BP-514 Battery
Be ready to capture that great shot - keep a spare battery on hand! Unlike NiCad batteries lithium ion batteries have no ''''memory effect'''' to reduce their capacity.--The BP-514 Battery Pack from Canon p . . . .
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Society for Human Resource Management
Employer Incentives for Hiring Individuals with Disabilities
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· OZON.ru 10320р. [Проверить наличие]
ISBN: 1586440438Издательство: Society For Human Resource Management
Год издания: 2004
Book DescriptionThis report presents the results of a survey conducted by the Society for Human Resource Management in conjunction with the Cerebral Palsy Research Foundation, University of Texas at San Antonio, and Wichita State University to gain an understanding of why low employment rates for individuals with disabilities persist. Presented is information on how knowledgeable human resource professionals are regarding various governmental incentives for hiring individuals with disabilities, how many companies actually take advantage of employer incentives, and who in companies make decisions about using them. The attitudes and opinions of human resource professionals regarding the impact of the Americans with Disabilities Act on the employment of individuals with disabilities are also discussed. Other topics covered include the level of effort companies expend in recruiting individuals with disabilities; senior managements' personal experiences with disabilities, and ideas from...
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Education, News & Politics, Society & CultureFollow
RN Breakfast is the program informed Australians wake up to. Start each day with comprehensive coverage and analysis of national and international events, and hear interviews with the people who matter today—along with those who'll be making news tomorrow.
April 22nd, 2015
Episode 620 of 2961 episodes
As Australians flock to Gallipoli for the centenary of the Anzac campaign, it's sobering to remember that the Turkish loss of life was twice that of the allied forces.
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Fast movement and great balance in this classy all-round scissor.
Made from hardened stainless steel alloyed with chrome for durability and great looks, these smashing little all-rounders will make an invaluable addition to your professional toolkit, with hollow-ground, slightly convex blades giving you great slice-cutting capability. They have a pivot point specially designed for really fast movement and great balance, so they feel really easy and comfortable to use for any kind of technique.
The black, titanium-coated finish looks great, protects your fingers form nickel allergies and gives the Slicy extra-long cutting performance.
The Tondeo Slicy comes with classic (even) handles. It's supplied with removable finger-ring inserts and an oil dropper.
Press fastening leather sheath that holds one pair of hairdressing scissors.
Contents shown not included
Look after your scissors with Kobe's scissor care kit including a scissor cloth, high quality scissor oil and 3 different sizes of finger-ring inserts.
Revitalise your tools with our professional services
Copyright © 2005-2017 CoolBlades.


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Urge your members of Congress to support continued medical research funding
Seven locations, one great program
The only official Highlights of ASH®. Learn how new clinical research presented during the preceding ASH® Annual Meeting will influence hematology patient care and what to expect in emerging drug therapies.
North AmericaAsia-PacificLatin America
September 8 - 9, 2017
Chicago, IL
Hear the top experts in hematologic malignancies discuss the latest developments in clinical care and find answers to your most challenging patient care questions.
December 9-12, 2017
(Exhibits: December 9-10, 2017)
Atlanta, Georgia
The American Society of Hematology (ASH) invites you to Atlanta, Georgia, for its 59th annual meeting. As the premier hematology event of the year, this meeting is an invaluable educational experience for all attendees.
Information from the 2016 ASH Annual Meeting
CME & Certificate of Attendance
2016 Abstracts Online
2016 ASH Annual Meeting Webcasts
Instantly access webcasts from ASH's many educational meetings and webinars.
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Zig Zag Zoom$29.99
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From HardDrive Labs: Make your Webcam capable of zooming, panning, motion detection, recording video, photos and upload time lapse photos to the internet so that all can view with Zig Zag Zoom. There are not motors necessary to pan and zoom when using this software. Add security to your Webcam with the adjustable motion detection feature. If anything moves in front of your Webcam this software will snap a picture of it so that you can view what your Webcam witnessed. You can view time-lapse photos over the internet allowing you to check up on the kids and your house. You can also use this feature to start your own Webcam site with a time-lapse view of anything you want to publish so that everybody can share the same view. Record movies on your Webcam by using the video record feature and take a picture by using the snapshot feature.
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Almost Nightfall
An older couple contemplates life on a sunny afternoon.
Emma nodded when the pond came into view, and John slowed the sedan onto the roadside and quieted the engine. They eased out and pulled a blanket and a brimming wicker basket from the back seat. They surveyed the pond. It was filled with geese today, the birds having also claimed the embankment. They moved to the water's edge and he spun the blanket outward. The geese scattered far enough to allow the cloth to drift to the ground in an almost perfect square. The valley's familiar diversions - the white steeple a mile away, and copses scattered acres apart - poked above the low brush. The afternoon heat was building, and John eased onto his elbows as he listened to the rustlings in the bramble, eventually falling asleep to the valley's natural rhythms.
He awoke to the rustling of paper. Emma had pulled a wrinkled bag from the basket. It was filled with the ends of bread loaves, which she tore apart, scattering the crumbs about her as she rose and circled the water. The bits of dough drew the attention of three goslings, and they waddled behind her nuzzling at the smallest pieces. Emma delighted in their odd parade until a large goose made chase. Honking, with her wings spread, the dithered fowl corralled her brood and marched them away. Emma turned, and laughed an undignified laugh before offering her apology for the intrusion.
Alone, John lowered his glasses which he had set across his brow, and placed them on the tip of his nose. From his back pocket he withdrew an envelope that bore an elegant script running its length. Inside, the typed sheets of the will were long, luxuriant, nearly the texture of parchment, and bound together by brass fasteners. He scrutinized the words, and as Emma returned, still bright from her encounter, he extended the pages to her. Her smile evaporated, and she turned away, shooing the elderly man with a wave. He motioned for her to sit next to him. Instead, Emma closed her eyes and tilted her head toward the sun. When she lifted her hat's brim to expose most of her lined face, he withdrew the pages and folded them away.
Again, she left their modest encampment. This time to walk a graveled path, her arms in a fold, her nape at a gentle arch. Emma wore thick-soled shoes for the outing, and she kicked and poked at the small rocks, often unearthing the stones from her path. Not until much later, when the air had cooled, and Emma had returned and wrapped herself in a thick sweater, and each of their shadows had distorted, did the couple relinquish the day.
The geese left singly throughout the afternoon and now none remained. John folded the blanket in halves, aligning the ends perfectly until it tucked neatly under his arm. His hand found the small of Emma's back and he guided her through the gloaming. It was almost nightfall when they drove through the valley, and the beams of light bounced unpredictably over their irregular road.


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Dresser - Fontanet - New Goshen - North Terre Haute - Riley - Seelyville - Shepardsville - St. Mary of the Woods - Tecumseh - Terre Haute - Toad Hop - West Terre Haute
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The Complete Guide to Solo Travel in Europe for Women — Tips for Having an Amazing Time While Traveling Alone
We've rounded up our 10 best carry-on bags for every traveler. Click the infographic to find out which bags made the cut and why. #travel #travelproducts #bags
To prevent bottles from leaking in your luggage, place plastic wrap over the container and then seal it with the cap. | 28 Brilliant Travel Hacks You Need To Know For Summer Vacations
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Best Travel Backpack For Women | Best Travel Backpacks for Women - Review of Designer Luggage Backpacks ...
The Top 10 Tips for Traveling Cheap and Keeping it Authentic in Germany #BudgetTravel #Germany
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Whether you're traveling for #business or for pleasure, you don't want to waste time at baggage claim. Check out these best carry-on bags! #Travel
«With love» from Burberry for Christmas ~ They can sent me a parcel & most women ...ANYTIME! Just get it to me soon! ~
Portable Traveling Dot Patterned Bra Bag Eva Bag Underwear Bag for Women Ladies from UltraBarato Gadgets
10 Stylish Camera Bags for Women. Well, I was not going to pin this post at first, because I thought the idea of a purse camera bag was lame, and would not work. I was wrong! After I saw the pictures of how organized, and how much camera gear you can fit
Large handbag in Vintage style. It has comfortable handles and wide compartment protected by flap. It is made from high quality leather and offers plenty of storage space. Perfect for work or weekend trips as necessary.
Weekenders are the perfect alternative to your large duffel, with all the same useful qualities and just the right amount of space.
Travelling With Kids Part 1- Carry-on Luggage for Air Travel - Our Little House in the Country, #travel #airtravel #travelwithkids
Large bag in retro style. It is made from high quality leather and has convenient handles. It includes wide compartment and side pockets. Great gift idea. Neutral design for man and woman.
Personalized label made of high quality leather. It can be used as a keychain or luggage label. Excellent gift idea. It has many positive recommendations from customers.
Bric's Bojola Collection Luggage. I can't actually find this bag on the linked website, but I like the white and brown leather combination. Maybe someone else makes a similar bag
MARC BY MARC JACOBS 'Globetrotter' Travel Wallet | womens wallet | womens style | womens fashion | womenswear | wantering
Tory Burch Monogram travel accessories: DO leave home with these… and give them to everyone you know. #EditorsWishlist
9 Free (or REALLY Cheap) things to do in London! in Advice, England, London, United Kingdom | Travel | Hand Luggage Only


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racheljurado1 on Aug 16, 2016
This is my second portrait from Magnetically Charged that I got for some friends as a present, and it was perfect! Special thanks to Alicia for finishing the portrait and even getting it in the mail the day after ordered it.
Megan peterson on Dec 20, 2015
Alicia is super talented!! She brought out my dogs features and their personalities in the painting and never even met them! She was quick to respond and I just love the painting! Thank you Alicia!!!
olivia whitehead on Dec 8, 2015
This is the second custom portrait I have purchased and I love it!
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I am a 20-something happily married graphic designer living in Upstate New York, originally from Oklahoma, named Alicia. I love Cheetos, Lord of the Rings, schnauzers, red lipstick, a good book, a good cuppa, vintage clothing, and cameras.
Shop policies
Accepted payment methods
- Accepts Etsy gift cards
If you ordered a custom portrait, it will take about 1 week to finish depending on where you are in the queue. And then will be shipped out as soon as you approve it!
Upgrades are also available if you want to use a Flat Rate cardboard envelope via USPS, or message me before purchase and I can make a custom listing if you prefer using UPS.
Refunds and Exchanges
Also, if you order a custom portrait and are unhappy with my painting or rendering before I ship it out, I will refund 50%+shipping or one "redo" painting. If you're not happy after the redo, you forfeit the partial refund. This is to account for expenses and time lost.
Additional policies and FAQs
Customs fees are the responsibility of the buyer. I ship out everything in person at the post office to ensure I pay postage correctly and complete all forms necessary.


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"Wednesday is looking like a pretty messy day" according to Jeff Coulson of Environment Canada. "It looks like we'll see some snow begin overnight which will mix into some ice pellets as we get further into the morning commute."
Environment Canada issued a special weather statement for the City Of Toronto on Wednesday, revealing that a major winter storm is expected to affect commutes on Wednesday and Thursday.
A low pressure system is emerging from Texas and is expected to intensify bringing significant amounts of rain, snow and ice pellets, along with the possibility of freezing rain.
"These types of winter storms from Texas typically bring 15 to 30 mm or more of rain" said Coulson "We could see snowfall and freezing rain warnings issued in the GTA, depending on the timing of the precipitation."
Travel conditions are expected to quickly deteriorate Wednesday morning as Coulson believes with the current forecast, the bulk of the morning commute should be slow for drivers.


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"The old trail" Åndalsnes Things to Do Tip by sim1
Åndalsnes Things to Do: 40 reviews and 41 photos
The 'Trollstigen' has always been an important connecting routes between Sunnmøre to Romsdal. The original route was not this road, but a narrow horse trail. You can still see this horse trail and the good news is that it is restored and is now used as a hiking trail. It sounds like a beautiful trail to hike, although probably very steep and exhausting to do.
If you do like me and take the car instead, you might have to think about which season to travel here. During the winter to approximately the middle of May the road is closed, depending on snow conditions.
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Stony Brook has a population of 13,727 and is located in Suffolk county.
For New York residents, the average tuition for all Stony Brook emergency management schools for the 2009 - 2010 school year was $4,970.00. For non-residents, the average tuition for Stony Brook emergency management schools was $12,870.00.
The cost, on average, for books and supplies needed for emergency management schools in Stony Brook is $900.00. The average cost for room and board for students living on campus at emergency management schools in Stony Brook was $9,590.00. The average cost for students living at home was $6,760.00 for the 2009 - 2010 school year. Students have a choice of one emergency management college in Stony Brook to attend. 30 students were reported to have graduated from emergency management programs in Stony Brook in the 2008 - 2009 school year.


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Participate, send your picture with your lovely MOM on this year mother's day with coolest caption, hashtag, and tag with friends and win exiting prizes.
Mothers Day Contest | Participate | Send Picture With Mother | Get Gifts
AsCoolAsMom Contest is issued on the facebook page of orient-electric on mother's dat. You know friends this week we celebrate Mothers day 2016. Just send your picture with your mom with caption and have a chance to win exiting prizes. For participate jus upload your picture with your MOM with the best caption or tell why your MOM is exiting prizes, Comment, tagging @orientelectric and use #AsCoolAsMom and win. The best coolest entry has a chance to win prize. If you are share your Mother's love then participate by on click link:-
Contest Ends: May 8th, 2016 at midnight
If you have any doubt and interested for get more information the go to Link:-


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Cadillac Dover DE
Locate a Cadillac car dealer in Dover, DE. Get a free quote on a new or used Cadillac or contact a Cadillac dealership in Dover for financing, incentives, and Special Offers.
Cadillac CRX Dover DE
This page provides relevant content and local businesses that can help with your search for information on Cadillac CRX. You will find informative articles about Cadillac CRX, including "The 2010 Cadillac SRX Turbo Premium". Below you will also find local businesses that may provide the products or services you are looking for. Please scroll down to find the local resources in Dover, DE that can help answer your questions about Cadillac CRX.


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iPhone Job by
- Status Closed
- Budget $10000 USD
- Total Bids 1
Project Description
Hello Beepig,
We are looking for your team to develop a mobile dating application on a fixed bid. The project will begin with the iPhone, then Android, then Blackberry and other areas including ipad and other devices. The app will be a sophisticated application utilizing GPS, Accelerometer, Camera, Facebook login and other key aspects. The application will also work against the Amazon cloud. The developer would handle all aspects of development from interface to backend and services.
Please let us know if such an application is something you feel comfortable developing? If so what your earliest availability to begin such project with us would be?
Please let me know, we are looking to begin this project shortly.
Thank you,
BenGet free quotes for a project like this
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Press / Media Relations Account Executive jobs in Travel and Tourism
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Senior Account Executive - Free Trip to The Caribbean!
Fancy telling your friends you're off to the Caribbean to work on a campaign? Have high-end luxury experience?
Senior Account Executive - Junior Account Manager - High End Luxury
Want to go to the Carribean? Want to work in a company that is creative and buzzy?
PR Account Executive role – Luxury Travel, Restaurant + Lifestyle
Our central London based client, is an independent creative comms agency, that oozes luxury. Specialising in travel/hotels, restaurants and lifesty...
Cut the London commute! Enjoy fantastic PR opportunities in Northern Home Counties
Passionate about PR but sick of the daily grind in central London? Awesome Senior Account Executive opportunities with Northern Home Counties based...
PR Account Executive
Fancy starting your travel PR career in an award-winning and dynamic agency?
Account Executive - Luxury Property PR / Corporate Communciations
INTERNS and GRADUATES welcome to apply! NO PREVIOUS EXPERIENCE in Property PR is necessary. For full info / spec, apply
Communications Executive - In-House
Can you start work Monday 27th February?
Account Manager - Director - We're Hiring!
We have a number of great opportunities for consultants who work within the b2b sector and or technology PR. We can offer you the chance to work fo...
Senior Account Executive, Luxury Brands, Central London
Rare opportunity for an ambitious Senior Account Executive to join a boutique Luxury Practice within a top global agency.
Senior Account Executive - Property PR
Work for an award winning large PR company who offers excellent benefits including gym membership and generous bonus.
Luxury lifestyle PR - Account Executive/Senior Account Executive
A luxury communications agency based in Chelsea is looking for a new Account Executive or Senior Account Executive to join their team. The agency s...
Junior roles in luxury lifestyle
This leading luxury lifestyle agency is looking to boost their PR team with juniors at JAE, AE and SAE level
Account Executive - Issues/Charity/Public Sector/Consumer
Wow what a role! One of our all time favourite clients is seeking a talented account executive to join them. They are an 80 strong integrated agenc...
Account Executive, Consumer Brands, Consumer and Corporate, Central London
Global agency seeks new superstar at Account Executive level!
Account Executive / Senior Account Executive – Luxury Lifestyle & Travel PR
An international lifestyle agency is looking for an energetic Account Executive or Senior Account Executive to join their dynamic team. Working acr...
Press Officer, Travel & Leisure, Hotel Chain, Oxfordshire
Press Officer role with an events edge. Rare in-house opportunity at a hugely successful hotel group!
Account Executive - Amazing Consumer Brands
We have two fantastic Account Executive roles in hot off the press. 1. A superb medium size agency based in Farringdon. They have a superb reputati...
Account Executive – Premium Drinks + Travel Brands
Passionate about drinks & travel? Awesome Account Executive required! Top tier lifestyle agency & an amazing client list featuring fantastic drin...
Account executive - Four Travel
Four Travel is looking for an enthusiastic, proactive and ambitious PR account executive to provide PR support.
Senior Account Executive - Issues/Public Sector/Charity
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Arab-American Heritage Month
MBN celebrates Arab-American Heritage Month
MBN commemorated Arab-American Heritage Month with special shows and acquired programming that highlights the rich contributions made by Arab-Americans to the U.S. culture. Alhurra's Al Youm traveled to Dearborn, Mich., to profile the city and community that is considered the unofficial capital of Arab-Americans. The show explored the millennial generation of Arab-Americans to see if they face the same challenges of earlier generations. From New York, the show examined the Yemeni community and how they are preserving heritage. Al Youm profiled the Center for Arabic Culture in Boston and its work to maintain cultural bridges between East and West. The show also examined the growing hijab business in the U.S. and profiled Arab-American comedians.
MBN Commemorates Arab-American Heritage Month
April was Arab-American Heritage Month and the Middle East Broadcasting Networks, Alhurra television and Radio Sawa commemorated the occasion with special news and programming highlighting the achievements of Arab-Americans, and their integration into the American society.


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Last Updated: 2/21/2017 6:34 PM
Description for 3225 Bob Wallace Avenue Huntsville, AL 35824
Madison County
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From Downtown take a right onto Bob Wallace travel 0.6 miles. Professional Center will be on your left.
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Huntsville Real Estate
Huntsville is located in Alabama. Huntsville, Alabama 35824 has a population of 179,855.
The median household income in Huntsville, Alabama 35824 is $48,632. The median household income for the surrounding county is $58,242 compared to the national median of $53,046. The median age of people living in Huntsville 35824 is 36.6 years.
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Week Twenty One As we travelled down the coast to our next stop, the well-known Freycinet National Park, Coles Bay and the base for the hike up to the Wineglass
10 DOGGY-DOS for what to pack Hi again! Campdog Maggie back to clue you in on what to pack when travelling with your best friends – dogs of course! We dogs
Happy Campers Series Episode 6 – The Mulcahy Family Grampers The Mulcahy family are longtime "grampers", or in other words, multi-generational campers. Mum and dad, nan and pop, and the kids
Are you planning a big caravan adventure? A lap around Australia perhaps? A trip down the coast to a remote beach or a family-friendly caravan park? No matter what trip
Week Twenty After loading up on supplies in St Helens we loaded the van with water and headed north 15km to our next camp in the Bay of Fires. The Bay
Week Nineteen One of the places we have always been most excited to see is the Bay of Fires on the north-east coast of Tasmania so we decided to pack
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Contact kmwordsmith
Phone: (585) 266-4818
Email: firstname.lastname@example.com
Phone: (585) 266-4818
Email: firstname.lastname@example.com
I'm currently working on projects with the following clients:
All my best ,
"Karen, was a pleasure to work with. We engaged her to write and edit web pages dedicated to health research and she did a superb job. She has a clear and concise writing style – perfect for the web – and was able to communicate at a level appropriate to our audience. Her research was thorough, the presentation of information was logical and accurate, and she delivered copy on deadline. I would welcome the opportunity to work with her again!"
Gwenn Voelckers, Director of Health Communications
University of Rochester Medical Center
"You really found the nuance in delivering the telehealth benefits for our annual appeal. I must say that I appreciate your work ethic and enjoy working with you all the more. You're one of 'my things I'm thankful for,' this Thanksgiving!"
Jean Pope Boyle, Director of Development
Seniorsfirst Communities & Services
"Karen Marley learned our project inside and out, conducted interviews with the project team, and created content that was well-researched and unique. Karen was very responsive to our short time schedules and delivered on time. In addition to always being friendly, Karen responded quickly to our inquiries and presented us with quality work. She has a strong command of the English language and uses it in appropriate and creative ways.
For anyone who wishes to put forth a polished and professional product, without hesitation I would recommend Karen. Kmwordsmith is the perfect content solution."
Anthony J. Costello, CEO
The Costello Group: U.S. Airports, Clinton Crossings Medical Center, The Reserve on the Erie Canal, CityGate
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Is Sunward Pharmaceutical incorrect or missing key business information? Update your Hotfrog listing today.
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Hotfrog Singapore provides information regarding Sunward Pharmaceutical in Singapore. Sunward Pharmaceutical is located at 11 Wan Lee Rd, Singapore 627943 and provides Healthcare services. Contact them on 6265 6022 or by visiting them on their website http://www.sunwardpharma.com.
Is Sunward Pharmaceutical in 11 Wan Lee Rd, Singapore 627943 your business? Claim your listing and attract more leads by adding more content, photos and other business details.
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Happy Weekend from Brooklyn Bridge!! Someone, please take me back!
NYC Morning At Brooklyn Bridge
Protect Your Eyes In A Stylish Way
Winter Wonderland with Free People
Styling a minimal yet stylish, fun winter look in a collaboration with Free People at one of the favorite places on the earth!


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'Undie Run' requires as little clothing as possible
College students strip down to their underwear to participate in the annual Undie Run in Tempe, Arizona. The run encourages students to donate the clothes off their backs to charity. VPC


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Edit Article
How to Make a Revision Timetable
As we are nearing the exam season, it is still not too late to start revising. Here are a few tips to making an effective revision timetable that will help you get the grades you deserve.
1Get some paper and some different coloured pens. A ruler might help, too, if you want to make a table rather than a list.
2List all the subjects you do and break them down into sub-categories. English Literature, for example, may be broken down into the different books you are studying and then further into chapters/themes/quotes etc. Highlight or circle the different subjects using different colours.
3Mix your subjects up. Don't put all your nightmare subjects on one day, and the ones you like on another day.
4Decide which tasks you should only spend short bursts of time on and you should maybe spend a whole day on to get out of the way. The second should be reserved for big projects that you won't get tired with.
5Allocate days for the big projects. Do this first, keeping in mind you may need to go back and change things later, so allow time for this.
6Out of the remaining days, make sure you write down any other commitments you have e.g. a job, lessons, meeting, or social events.
7Decide what times you work best and put the tasks that require more energy or concentration during your optimum times.
8When putting the rest of your revision into your diary or table, put times down and make sure you start each day reasonably early.
9Start your day off with something you know you can accomplish. This will give you more motivation for the rest of the day. Make sure the next slot is something heavier so you don't put off your worst subjects until the end.
10It has been said the optimum concentration time is 32 minutes so keep this in mind when deciding how long each session should last.
11Give yourself regular breaks. If when doing your revision you are doing well, you can always decide to keep going.
12Cross things off as you do them because it will give you a sense of accomplishment.
13The most important thing - don't panic. In addition, make sure you get lots of sleep before the exam day.
Community Q&A
What is the best time to study?wikiHow ContributorThat is largely up to you. Think about when you have free time in your schedule, and try to pick a time when you'll be awake, alert, and free of distractions.
How do I maintain a positive outlook?wikiHow ContributorMake a "to do" list and have a planner so that you know what you need to be doing. When a person is organised, he/she will automatically be more positive. Congratulate yourself on your accomplishments and if there's something you can't get done, don't be hard on yourself.
Ask a Question
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If this question (or a similar one) is answered twice in this section, please click here to let us know.
- Be realistic about what you can achieve but remember it is all about trial and error. Even if it goes wrong, you can always adjust your timetable.
- When you are drawing your timetable, draw one with just the subjects you have to revise (e.g. math, chemistry) and then draw another one with which topics within each subject to revise (e.g. angles, periodic table). This way you have a quick table and one, which is more precise, detailed, and focused!
- Studying with a friend can be productive and fun. You can revise subjects you both do and test each other on things you've learned. Even if you don't do the same subjects, you can motivate each other to work and help each other stick to your revision plans.
- Make yourself a detailed schedule the night before - especially if it's a weekend. For instance, 9:30 - Get up, take a shower. 10:00 - eat breakfast, grab some coffee etc. 10:30 - Revise the Russian Revolution... and so on! You'll be surprised at how much more likely you are to get work done if you have everything planned out exactly.
- Don't spend ages making your timetable look pretty. If you are a perfectionist and having a messy timetable will unnerve you, make it look clean and tidy, but don't spend precious revision time on drawing pretty pictures or colouring things in.
Things You'll Need
- Paper or a diary
- A Pen
- Coloured pens
- Ruler
Article Info
Categories: Calendars Planners and Schedules
In other languages:
Español: elaborar un horario de repaso, Italiano: Programmare un Ripasso Scolastico, Português: Organizar uma Planilha para Revisão de Estudos, Русский: составить график по подготовке к экзаменам, Deutsch: Einen Lernplan basteln
Thanks to all authors for creating a page that has been read 173,647 times.
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New Mansion House
Real Reviews
Real reviews from fellow travellers
These are trusted reviews from guests who have actually stayed here. We hope you find them helpful.
Note: we only give a property a rating once it has received a minimum of 3 reviews.
- Page:1/1
- " Good location, however, I wouldn't recommend staying in this one - my shared room was too small, there is no lift so you have to carry your luggage on narrow stairs if you stay on top. The room was overcrowded, occupied with long term residents, who had a weird behaviour to eat at 00:30 and talk, cheap breakfast without butter and just one jam (strawberry), a poor facilities, old building that needs renovations. Some hostels twice cheaper have a better value than this one. "
- 13 Sep 2016 Anonymous Lithuania
- " Nice clean place to stay. It is a hostel where people stay for a longer time so there are no hostel activities like a pub crawl. But it has big clean rooms and the bathroom are clean as well. no elevator but a lot of stairs.The breakfast is early 7:30 and only a hour long. Otherwise everything you need including wifi everywhere. "
- 27 Jun 2016 Laura Germany
- " I haven't got thoughts about the experience ,only my option that the room very clean .toilet and shower facility clean and breakfast provided. "
- 23 Feb 2016 miss tania ann British Virgin Islands


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Not a Member? Get access to HR news and resources that you can trust.
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Nearly one-half of organizations (46%) use virtual teams in their workplace. Roughly one-quarter of the organizations using virtual teams are U.S.-based operations (28%), while organizations with multinational operations are more than twice as likely (66%) to use virtual teams. Approximately one-half of organizations using virtual teams use them to include talent in different geographic locations (53%) and to boost collaboration among employees in different geographic locations (49%). Brainstorming solutions for problems or issues (72%) was reported as the most successful behavior for organizations using virtual teams.
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